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HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

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UNIVERSITY OF KANSAS

SCIENCE BULLETIN

APR 2 11960

XJNIVERSITY OF KANSAS PUBUCATIONS

University of Kansas Science Bulletin - Vol. XL

April 20, 1960

Lawrence, Kansas

ANNOUNCEMENT

The University of Kansas Science Bulletin (continuation of the
Kansas University Quarterly) is issued in parts at irregular inter-
vals. Each volume contains from 500 to 1,800 pages of reading
matter, with necessary illustrations. Exchanges with other institu-
tions and learned societies everywhere are solicited. All exchanges
should be addressed to

The UNivERsrrY of Kansas Science Bulletin,
Library of the UNxvERsmr of Ka.nsas,

Lawrence, Kan.

PUBLICATION DATES

The actual date of publication (i. e., mailing date) of many of the
volumes of the University of Kansas Science Bulletin differs so
markedly from the dates borne on the covers of the pubHcation or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
time as the whole volume.

VoL

XX— October 1, 1932.
XXI— November 27, 1934.
XXII— November 15, 1935.
XXIII— August 15, 1936.
XXIV— February 16, 1938.
XXV— July 10, 1939.
XXVI— November 27, 1940.
XXVII, Pt I— Dec. 30, 1941.
XXVIII, Pt I— May 15, 1942.
Pt. II— Nov. 12, 1942.
XXIX, Pt I— July 15, 1943.
Pt II— Oct. 15, 1943.
XXX, Pt I— June 12, 1944.
Pt II— June 15, 1945.
XXXI, Pt I— May 1, 1946.
Pt II— Nov. 1, 1947.

Vol.
XXXII— Nov. 25, 1948.
XXXni, Pt. I— April 20, 1949.

Pt II— March 20, 1950.
XXXIV, Pt I— Oct 1, 1951.

Pt II— Feb. 15, 1952.
XXXV, Pt I— July 1, 1952.

Pt II— Sept 10, 1953.

Pt III— Nov. 20, 1953.
XXXVI, Pt I— June 1, 1954.

Pt II— July 15, 1954.
XXXVII, Pt. I— October 15, 1955.

Pt II— June 29, 1956.
XXXVni, Pt. I— Dec. 20, 1956

Pt. II— March 2, 1958
XXXIX— November 18, 1958

Editor Edward H. Taylor

Editorial Board . .

R. H. Thompson, Chairman

Charles Michener

Paul Roofe

David Paretsky

Worthie H. Horr

Parke H. Woodard, Secretary

UNIVERSITY OF KANSAS

SCIENCE BULLETIN

nEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

Volume XL

University of Kansas Publications

Lawrence, April 20, 1960

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THE STATE PRINTING PLANT

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Contents of Volume XL

Number

1. Fragmentary Remains of a Lycopod from the Desmoines-

ian Series (Pennsylvanian) of Southeastern Kansas,

Arthur Cridhind,

2. A New Softshell Turtle (Genus Trionyx) from Coahuila,

Mexico Robert G. Webb and John M. Le^Ier,

3. A New CaeciHan Genus in India Edward H. Taylor,

4. On the CaeciHan Species Ichthyophis monochrous and

Ichthyophis glutinosus and Related Species,

Edward H. Taylor,

5. A Review of the Genus Paroxystoglossa (Hymenoptera:

Halictidae) /. S. Moure, C. M. F.,

6. Comparative External Moq^hology and Taxonomy of

Nymphs of the Trombiculidae (Acarina),

D. A. Crossley, Jr.,

7. Concerning the Genus Ventidius and Five New Species

( Hetoroptera, Gerridae ),

Herbert B. Hungerford and Ryuichi Matsuda,

8. A Revision of the Mite Family Rdellidae in North and Cen-

tral America ( Acarina, Prostigmata ) . . Warren T. Atyeo,

PAGE

21
31

37
121

135

323
345

MUS. COMP. ZOOL
LIBRARY

APR 2 11960

HARVARD
UNIVERSITY

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 1

Fragmentary Remains of a Lycopod from the Desmoinesian
Series ( Pennsylvanian ) of Southeastern Kansas

Arthur Cridland

Abstract: Lepidodendron inoorei sp. nov. is described from six specimens
found in Kansas coal balls. The assignment of this lycopod to Lepidodendron
is arbitrary as there are no leaf bases on any of the specimens.

INTRODUCTION

Six coal balls have been collected from southeastern Kansas each
of which contain a fragmentary specimen of a distinct fossil lyco-
pod (figures 13-18). The specimens are all considerably crushed
and have been invaded by small Cordaites roots (Amyelon radicans
Williamson). Their fragmentary nature is such that if only one
specimen was available I would hesitate to describe it as new. As
none of the six specimens show any transition to known species of
lycopods I believe they are worth describing as a new species. The
poor preservation may indicate that these fossils were carried some
distance (presumably by water) before preservation. The speci-
mens were studied using the cellulose acetate peel technique de-
scribed by Joy et al. ( 1956 ) .

Lepidodendron jnoorei sp. nov.

Holotypc specimen: No. 20 in the Kansas Uni\ersity coal ball
collection, collected by Dr. R. W. Baxter.

Paratype s^jecimens: No. 826 and No. 831 in the Kansas Univer-
sity coal ball colection, collected by Dr. R. W. Baxter. No. 1078 in
the Kansas University coal ball collection, collected by Dr. W. G.
Chaloner. No. 1043 in the Henry Shaw School of Botany coal ball
collection at Washington University, St. Louis, Missouri, collected
by Mr. T. Phillips in July 1958.

(3)

4 The University Science Bulletin

Locality and Iwrizon: All of the specimens were collected from
the Pittsburg and Midway Coal Company's strip mine north of
Hallowell, Cherokee County, Kansas. ( Section 4, T. 33 S., R. 22 E. )
The coal balls are from the Mineral or Fleming Coal of the Des-
moinesian Series. A full classification of these strata is given by
Searight et aJ. (1953).

Associated nuitcriah In fixe of the coal lialls ( Nos. 826, 831,
1020, 1043 and 1078) parts of Cordaitcs are commonest, particularly
the leaves. Other fossils present are Cordaitcs stems, Cordaitanthiis
cf. concinmis (Delevoryas) Fry, Cardiocarptis spinatus Graham,
Aletlwptcris sp., Myeloxylon sp., Lepidodcndron leaves, a Lepido-
carpon cone, a small twig of Lepidodcndron serratum Felix, Stipi-
toptcris cf. f^rac;"//.? Morgan and Delevoryas and fragments of a
Dolcrothcca. In the sixth coal ball (No. 20) the plant remains are
mainly Scolecopteris and large Psaroniiis roots. Other species pres-
ent are Amyclon radicans Williamson, Botryopteris petioles and an
undescribed pteridosperm stem.

Dia<inosis: Cortex of thick-walled cells, with distinct, parallel,
vertical strands of slightly thicker-walled cells. Metaxylem sca-
lariform, bars with a tendency to bifurcate and to become reticu-
late. No fine \ertical striations between the scalariform bars. No
corona. No secondary wood. Areas of decay around leaf traces
regularly arranged in both vertical and diagonal series.

The species is named after Mr. Kenneth Moore of the Pittsburg
and Midway Coal Company through whose co-operation it is
possible to collect coal balls at the strip mine north of Hallowell.

Description of the holotype. This specimen is an unbranched
crushed stem which is 10 cm. long and 74 mm. x 7 mm. in transxerse
section. There are 21 strands of thick-walled cortical cells sur-
rounding the exarch siphonostele (fig. 15 and 19). The stem
runs into the coal at the edge of the coal ball. In life it was slightly
wider than the measurements given and had one to several more
strands of thick-walled cells. The strands are mainly isolated from
one another but in areas where the preservation is better the strands
are joined to each other by intervening slightK' thinner-walled
cells (fig. 4 and 11). In transverse section the strands are
round and about 2.3 mm. in diameter or pear-shaped and about
2.5 mm. X 3.5 mm. (fig. 19). In the outer part of the strands
there are localized areas of small cells which grade in size to the
surrounding larger cells. These areas of small cells are frequently
partly or extensively decayed and the area of decay may extend
into the surrounding larger cells. The region occupied by the

Fragmentary Remains of a Lycopod 5

smaller cells is about 500 [j. across and the cells are polygonal or
roundish, 45 [x-60 \i in width. The surrounding larger cells are
similar in shape, about 180 [j. across, or they may be elongated radi-
ally and about 200 p. x 150 tj.. In cells least decomposed before
fossilization the walls may reach 11 [a in thickness from cell lumen
to cell lumen. The cells joining the strands to each other are also
thick-walled but their walls are thinner (about 7 [), from cell lumen
to cell lumen) than the walls of the cells in the main strands. Many
of the cortical cells are poorly preserved. Parts of their walls have
separated and at first glance the cell walls appear to be very thick
(fig. 21). By comparison with the sequence of cell-wall degrada-
tion established by Barghoorn ( 1952 ) I presume that these are rem-
nants of the outer part of the secondary wall and a dark residue
between them is all that remains of the inner layers of the secondary
wall and the primary wall. Tangential sections show that the strands
of cortical cells are parallel and in areas of better preservation
thinner-walled cells join the strands to each other (fig. 4). The
tangential sections were cut obliquely and although the length of
the thick-walled cortical cells could not be measured accurateb
they are more than 900 a long.

The exarch siphonostele is 7 mm. x 1.2 mm. from protoxylem to
protoxylem and is composed entirely of tracheids. There is no
definite corona but there are occasionally very small groups of pro-
toxylem cells projecting from the main protoxylem of the stele
{ figure 2 ) . These groups of cells are few, poorly preser\'ed and
ha\'e no regular arrangement. The main part of the protoxylem
forms a narrow zone at the periphery of the siphonostele. In trans-
verse section the cells of the protoxylem are 30 [;. in diameter and
the metaxylem tracheids are 120 ij. radially x 75 [x tangentially. The
walls of the tracheids are well preserved and have not decomposed
like those of the cortical cells. (The detailed structure of the tra-
cheids is the same as in .specimen No. 831 where the stele is de-
scribed in greatest detail). No leaf traces are to be seen leaving the
stele. There is no secondary wood.

Description of the poratypes. Specimen No. 831. This specimen
is similar to No. 20 but it is smaller. The specimen lies near the
edge of the coal ball and as a result part is missing. The part pre-
.served is a little more than 5 cm. long and is 19 mm. x 4.5 mm. in
transverse section. There are twenty strands of thick-walled corti-
cal cells still present around the stele (fig. 5 and 17). In trans-
verse section these strands are rounded (about 1.2 mm. in diameter)
to ovoid (about 1.3 mm. x 1.0 mm.). The cells joining the strands

6 The University Science Bulletin

to each other are not well preserved in this specimen. The thick-
walled cells of the strands are similar in shape and preservation
to those in specimen No. 20. They are from 48 [jl-120 [l in diameter
or else they are elongated radially and about 120 \). x 80 •^.. There
is no localized region of small cells in the cortical strands as there is
in specimen No. 20. Specimen 826 is the only other specimen lack-
ing this localization of smaller cells in the cortical strands.

In transverse section the stele is 6 mm. x 1 mm. from protoxylem
to protoxylem. There are small projections from the main protoxy-
lem of the stele as in specimen No. 20 but they are not numerous
and not conspicuous enough to form a corona. The metaxylem
tracheids measure 130 \i. radially x 67 t;. tangentially in transverse
section and they have scalariform tliickening on the radial and tan-
gential walls. The scalariform bars often bifurcate (fig. 8) and
frequently fairly robust vertical or almost vertical bars develop and
connect the scalariform bars giving a reticulate appearance (fig.
3). There are none of the fine vertical striations between the
scalariform bars which have been observed in so many fossil lyco-
pods (see Barghoorn and Scott, 1958). The scalariform bars of
the metaxylem tracheids are about 5 t;. in diameter and there is a
distance of about 10 [j. between the bars. The length of the tra-
cheids was not determined. The protoxylem consists of tracheids
which are about 30 \). in diameter and have spiral or annular thick-
ening (fig. 20). The thickenings are about 2.5 pL in diameter.

Specimen No. 826. This specimen is similar to specimen No. 831
but here the whole width of the stem is preserved. It is about
9 cm. long and 28 mm. x 3.2 mm. in transverse section. There are
25 strands of thick-walled cortical cells around the siphonostele
(figs. 7 and 13). These strands are 1.5 mm. to 2 mm. in diameter.
Tangential sections show that some slightly thinner-walled cells
are present joining the strands to each other (fig. 6) but they are not
abundant. The tangential sections prepared were oblique and
although the length of the thick-walled cortical cells could not be
measured accurately, they are more than 1 mm. long.

The crushed stele is 6 mm. x 1 mm. from protoxylem to protoxy-
lem. Both the metaxylem and the protoxylem are similar to that
described in specimen No. 831.

Specimen No. 1078. This specimen is at least 5 cm. long (only
one half of the coal ball was available for study ) and 40 mm. x 8 mm.
in transverse section. Twenty strands of thick-walled cortical cells
are preserved around the siphonostele (fig. 14). The strands of

Fragmentary Remains of a Lycopod 7

thick-walled cells are similar to those described in specimen No. 20.
Many of the cells joining the strands to each other are preserved
(fig. 10).

The stele is 6 mm. x 1 mm. from protoxylem to protoxylem in
transverse section and is of the same structure as those in the
previously described specimens.

Transverse sections of this specimen show a zone of thin-walled
cells outside the strands of thick-walled cortical cells (fig. 10).
This zone is poorly preserved but in several places it is clear that
these cells are joined to the thick-walled cells of the strands (fig. 1).
These thin-walled cells are squarish to irregular and about 110 [ji,
across. Radial sections show that this thin-walled tissue is con-
siderably broken.

Specimen No. 1020. This specimen is 11 cm. long and 72 mm. x
7.5 mm. in transverse section. It is heavily pyritized. Transverse
sections show 22 strands of thick-walled cells similar to those in
specimens No. 20 and No. 1078. These strands are completely
joined to each other by intervening cells (figs. 12 and 18). Tan-
gential sections of the cortex show that it has gaps in the tissue be-
tween the strands (fig. 23). These gaps are arranged in regular
vertical and diagonal series. They occasionally contain fragments of
tracheids having scalariform thickening which are interpreted as
the remains of leaf traces. The gaps in the cortex represent areas
of decay around them. The tracheids in these leaf traces are not
well preserxed but scalariform bars can be seen. The leaf traces
could not be followed back to the stele which is very poorly pre-
served.

The approximate size and position of the stele is indicated in
fig. 12. The few cells preserved show that it was an exarch sipho-
nostele. The metaxylem tracheids are about 100 jx radially x 75 \l
tangentially in transverse section. As the stele is badly decomposed
it is not possible to say whether these were the largest tracheids in
the stele. Scalariform thickenings were seen on the radial and ta-
gential walls of the metaxylem tracheids.

Speciynen No. 1043. This specimen is in a pyritized coal ball.
It is of interest because leaf traces are preserved. The specimen is
about 12 cm. long and 37 mm. x 7 mm. in transverse section. There
are 31 strands of thick-walled cells surrounding the siphonostele
(figs. 9 and 16). This number is approximate because all of the
strands are not completely isolated from each other. The strands of
thick-walled cells are ovoid and are 3 mm. radiallv x 1.8 mm. tan-

8 The University Science Bulletin

gentially in transverse section. The incli\ idual cells are about 100 [k
in diameter to 156 \t. radially x 108 ]j. tangentially. As in the other
specimens the cells of the cortical strands are considerably decom-
posed (fig. 24). In tangential sections the parallel nature of the
strands can be seen very clearly (fig. 22). The individual cells
are more than 800 \i long. The strands are joined into a cylinder by
intervening cells which are very well preserved in this specimen.
Large, regularly arranged gaps are present in the tissue between
the strands and most of these gaps have a leaf trace at one end ( fig.
25). The leaf traces in these gaps are not well preserved and arc
usually cut obliquely. Scalariform thickening can be seen on the
tracheid walls and in some cases fine vertical striations are present
between the scalariform bars but these are not very distinct. Where
the leaf traces in these gaps have been cut transversely they are
semicircular and are about 500 j. deep and 900 pi wide. The in-
dividual cells which are preserved are squarish and 180 ix across.
The cells near the center of the traces are poorly preserved but
there is a suggestion that the leaf traces are mesarch.

The stele itself is much broken but is clearly exarch. The esti-
mated measurement before fragmentation is 9 mm. x 1.2 mm. from
protoxylem to protoxylem. The metaxylem is composed of radially
elongated cells which are 120 \i x 62 \i. The protoxylem is also
elongated radially and the individual cells are 58 \). x 38 (j.. Unfor-
tunately the stele is surrounded by pyrite and although the preser-
vation of the outer regions is better than in the other specimens it
is still not as good as one desires. There are projections from the
outer edge of the stele. These are all interpreted as leaf traces.
Before the leaf traces separate from the stele they are elongated
tangentially and are 144 [x x 72 \). (fig. 24). When they are com-
pletely detached from the stele they are similar in shape but
slightly longer in the tangentially direction (160 \i \70 \).). There
are no obvious pointed projections from the stele which could be
interpreted as parts of a typical lycopod corona. The metaxylem
tracheids of the stele have scalariform thickening on the radial and
tangential walls and the scalariform bars occasionally bifurcate.
There are no fine \ertical striations between the scalariform bars.
The protoxylem tracheids have spiral or annular thickening. There
is no secondary wood.

DISCUSSION

The crushed stems vary in size considerably (figs. 5, 7 and 9-12).
The smallest specimen is 28 mm. x 3.2 mm. in transverse section
while the largest is 72 mm. x 7.5 mm. in transverse section. The

Fragmentary Remains of a Lycopod 9

stele is less variable in size and ranges from 6 mm. x 1 mm. to 9 mm.
X 1.2 mm. in transverse section. The general arrangement and
appearance of ihe tissues is the same in all the specimens and it is
safe to assume that they all belong to the same species.

The exarch siphonostele composed entirely of tracheids and the
scalariform thickening of the metaxylem suggest that the fossils
described here are lycopod stems. This is supported by specimens
No. 1020 and No. 1043 in which there is evidence that they bore
closely set leaf bases which were regularly arranged both in vertical
and diagonal series (figs. 22 and 23). The outer edge of the stele
does not show a corona, a structure which is characteristic of
known petrified lycopods, but its absence may be a result of poor
preservation. Very small, irregularly arranged projections from the
protoxylem of the stele in some specimens suggest that a corona
could have been present. No fine vertical striations are to be ob-
served between the scalariform bars of the metaxylem even under
examination with an oil-immersion objective. In some of the leaf
traces seen in specimen No. 1043 there are a few striations but they
are not as abundant as they appear even in poorly preserved speci-
mens of other fossil lycopods. There is no secondary thickening
even in the largest specimens.

The absence of leaf bases makes it impossible to determine to
which genus of lycopods the species belongs. The arrangement
of the leaf traces in vertical as well as diagonal series ( in contrast to
an arrangement in diagonal series only) is commonest in the genus
Sigillaria but is also found in LepidodencJron volkmanianiiin Stern-
berg. Rather than definitely assigning these fossils to Si'^illaria
without knowing the characters of the leaf cushions I have fol-
lowed Arnold ( 1940) for Lepidodendron johnsonii and Felix ( 1952)
for Lepidodendron kansanum in assigning fossil lycopods without
leaf bases to the genus Lepidodendron.

Several comparisons can be made with previously described
species of Lepidodendron. The figures of L. va.scidare Binney in
Leclercq (1925) are instructive. In Leclercq's plate 16 the mid-
dle cortex of the specimen has broken down into islands of cells
similar to the strands of thick-walled cells in L. moorei. This is
significant as it shows that similar decomposition of the cortex has
been recorded in at least one other fossil lycopod. There is no evi-
dence to refer the specimens described here to L. vasculare.

Of the lycopods described by Felix (1952) from southeastern
Kansas closest comparison can be made with Lej)ido(lendron di-
centrieum. This has a middle corte.\ which looks similar to the

10 The University Science Bulletin

strands of thick-walled cells of L. luoorei. In L. diccntricinn the
primary stele is wider than in L. inoorei. Also similar-sized speci-
mens show that whereas there is considerable secondary thickening
in L. clicentricittn there is none in L. moorei. Furthermore L. di-
centriciini has its primary xylem di\ ided into an inner and an outer
region. Nothing comparable to this has been observed in L.
moorei.

Lepidodendron serrafinn Felix from the same locality is a sipho-
nostelic form which like L. moorei has no secondary wood. There
is nothing in the structure of L. scrratum to suggest that on de-
composition strands of thick-walled cells similar to those in L.
moorei will be formed. The leaf cushions in L. serrotum cannot
be arranged in a vertical series as well as a diagonal series and its
metaxylem tracheids are larger in transverse section than those of
in L. moorei.

ACKNOWLEDGMENTS

I wish to express my thanks to Dr. R. W. Baxter for his helpful
advice and criticism during this work.

LITERATURE CITED

Arnold, C. A.

1940. Lepidodendron johnsonii, sp. now, from tlir Lowtr Pennsylvanian
of central Colorado. Contrib. Michigan Univ. Mus. I'aleont., vol.
6, pp. 21-52.
Barghoorn, E. S.

1952. Degradation of plant tissues in organic sediments. Journ. Sed.
Petrol., vol. 22, pp. 34-41, pis. 1 and 2.
Barchoorn, E. S., and Scott, R. A.

1958. Degradation of the plant cell wall and its relation to certain tra-
cheary features of the Lepidodendrales. Amer. Journ. Bot., \ ol. 45,
pp. 222-227.
Felix, C. J.

1952. A study of the arborescent lycopods of Southeastern Kansas. Ann.
Missouri Bot. Card., \o\. 39, pp. 263-289.

Joy, K. W. et al.

1956. A rapid cellulose peel techniciue in palaeobotany. Ann. Bot., N. S.
vol. 20, pp. 635-637.
Leclercq, S.

1925. Les coal balls de la couche Bouxharmont des Charbonnages de
Werister. Mem. Soc. Geol. Belgique, 1925, pp. 1-79, pis. 1-49.
Searight, W. V. et al.

1953. Classification of Desmoinesian (Pennsylvanian) of Northern Mid-
Continent. Bull. Amer. Assoc. Petrol. Geol., vol. 37, pp. 2747-2749.

Fragmentary Remains of a Lycopod

Figs. 1-4. hepidodendron moorei sp. nov.

Fig. 1 . Transverse section of specimen No. 1078 showing part of the thin-
walled corte.x (A) joined to a thick-wtilled cortical strand (B). Slide No.
213, X 100.

Fig. 2. 'i'ransverse section of the outer edge of the stele of specimen No. 20,
showing a small protoxx leni projection at the top of the figure. Slide .201,
X 400.

Fig. o. Fart ot a metaxyleni traeheid showing the tendene> h)r tlic sealaii-
lorin hars to hecome reticulate. Radial section of specimen No. 831. Slide
No. 207, X 400 (mounted sideways).

Fig. 4. Tangential section of specimen No. 20 showing the \ertical strands
of tliick-walled cortical cells. Some of the cells joining the strands to each
other are still present (shown hy diagonal lines). Slide No. 206, X 2 (mounted
sideways).

12 The University Science Bulletin

Fig. 5. Transverse section of specimen No. 831 showing 20 strands of
thick-walled cortical cells around tlic siplionostele. Slide No. 208, X 2.

Fig. 6. Tangential section of specimen No. 826 showing the vertical strands
of thick-walled cortical cells. The small amount of tissue joining the strands
to each other is shown by diagonal lines. SHde No. 211, X 2.

Fig. 7. Transverse section of specimen No. 826 showing 25 strands of thick-
walled cortical cells around the siphonostele. Shde No. 210, X 2.

Fig. 8. Part of a tracheid from specimen No. 831 in tangential section. Two
of the scalariform bars show bifurcations. SHde No. 209, X 400.

Fig. 9. Transverse section of specimen No. 1043 showing 32 strands of
thick-walled cortical cells around a broken siphonostele. The tissue joining the
strands to each other is shown by diagonal lines. SHde 291, X 2.

Fig. 10. Transverse section of specimen No. 1078 showing 21 strands of
thick-walled cortical cells around the siphonostele. The tissue joining the
strands to each other is shown by diagonal lines. The broken line represents
the limit of a thin-walled cortical tissue outside the strands of thick-walled cells.

Fig. 11. Transverse section of specimen No. 20 showing 20 strands of thick-
walled cortical cells surrounding the siphonostele. The small amount of tissue
joining the strands to each other is shown by diagonal lines. Slide No.
201, X 2.

Fig. 12. Transverse section of specimen No. 1020 showing 24 strands of
thick-walled cortical cells and a large amount of tissue joining the strands to
each other (shown by diagonal Hues). The position of the poorly preserved
stele is shown by stippHng. Slide No. 202, X 2.

Fragmentary Remains of a Lycopod

Figs. 5-12. Lepidodendron moorei sp. nov.

14 The University Science Bulletin

Figs. 13-18. Lepidodendrm moorei sp. nov. Transverse sections of the six
specimens showing the arrangement of the strands of thiek-walled cells around
the stele.

Fig. 13. Specimen No. 826. Slide No. 210, X 3.

Fig. 14. Specimen No. 1078. SHde No. 220, X 2.

Fig. 15. Specimen No. 20. Slide No. 201, X 2.

Fig. 16. Specimen No. 1043. SUde No. 291, X 2.

Fig. 17. Specimen No. 831. This photograph has been retouched to show
the position of the stele and the cortical strands. Slide No. 208, X 3.

Fig. 18. Specimen No. 1020. The stele is very poorly preserved in tliis
specimen. Slide No. 222, X 2.

Fracmentaky Remains of a Lycopod

Figs. 13-18. Lepidodendron moorei sp. nov.

16 The University Science Bulletin

Fig. 19. Transverse section of specimen No. 20 sliowing the stele and
surrounding strands of thick-walled cortical cells. Localized areas of sniall
cells can be seen in the two strands to the right-hand side of the figure. Slide

No. 201, X 8.

Fig. 20. Oblique tangential section through the protoxylcni of specmien
No. 831 showing the spiral and annular thickening of the tracheids. Slides

No. 209, X 500.

Fig. 21. Transverse section of the partly decomposed thick- walled cortical
cells of specimen No. 20. Slide No. 201, X 200.

Fragmentary Remains of a LvcoroD

■ av

Figs. 19-21. LepidodeTuIroii nioorci sp. nov.

18 The University Science Bullet

Fig. 22. Tangential section of the cortex of specimen No. 1043 showing the
parallel strands of thick-walled cortical cells which are joined to each other
by a slightly thinner-walled tissue. This thinner-walled tissue has regularly
arranged gaps. SHde No. 279, X 2.

Fig. 23. Tangential section of the cortex of specimen No. 1020 showing the
parallel strands of thick-wal'ed cells which are joined to each other by a
slighth- thinner-walled tissue. This thinner-walled tissue has regularly ar-
ranged gaps. The photograph is mounted obliquely. Slide No. 230, X 2.

Fig. 24. Transverse section of specimen No. 1043. The left-hand side of
the photograph shows the outer edge of the exarch siphonostele with a leaf
trace which is not yet detached. To the right of this there is a leaf trace which
is completely detached from the stele and is traversing the cortex. The lower
right-hand corner of the photograph shows some partly decomposed cells be-
longing to one of the strands of thick-walled cortical cells. Slide No. 263
X 100.

Fig. 25. An enlarged portion of the specimen shown in figure 22 showing a
leaf trace (T) at the upper edge of each of the gaps. The photograph is
mounted on its side so that the upper edges of the gaps are at the right. SHde
No. 279, X 5.

Fragmentary Remains of a Lycopod

Figs. 22-25. Lepidodendron moorci sp. nov.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 2

A New Softshell Turtle (Genus Trionyx) from
Coahuila, Mexico

Robert G. Webb and John M. Legler

For ten days in September, 1958, a field party from the University
of Kansas Museum of Natural History (KU) collected aquatic
vertebrates in central Coahuila, Mexico. Among the specimens are
14 examples of Trionyx obtained in the basin of Cuatro Cienegas.
Examination of these turtles revealed the presence of two distinct
species. Five of the specimens are Trionyx spinifer enioryi. The
remaining nine specimens and one other individual, in the Univer-
sity of Illinois Museum of Natural History (lU), represent a pre-
viously unrecognized species that, in allusion to its over-all blackish
coloration, may be known as:

Trionyx ater* sp. nov.

Black Softshell

?Amydu tnuticu, Miiller, Verb. Naturf. Ges. Basel, vol. 6, 1878,
p. 641.

Holotype. — KU 46903, alcoholic; female; obtained 16 kilometers
south of Cuatro Cienegas, Coahuila, September 6, 1958, by John M.
Legler, Wendell L. Minckley, and Robert B. Wimmer; original
number, JML 1708 (PI. I).

Paratypes. — A total of nine alcoholic specimens: KU 46904-6,
46908-10, 46912 (females), KU 46911 (male); same data as holo-
type; obtained September 6 to 8, 1958; lU 43510 (female), ob-
tained 5.7 miles west of Cuatro Cionegas, Coahuila, July 10, 1958,
by Pete S. Chrapliwy and Kenneth L. Williams.

Dicifinosis. — A species of softshell tintle most closely allied to
Trionyx spinifer enioryi and having: (1) uniform blackish colora-

* ater ( Latin ) — black.

(21)

22 The University Science Bulletin

tion (rather than a pattern of pale and dark markings) on the
carapace and dorsal surfaces of limbs, neck, and head; (2) ventral
surfaces heavily speckled with black; ( 3 ) no evidence of a pale mar-
ginal band on carapace (females only); (4) longitudinal corruga-
tions on the posterior part of the carapace (most females); (5) no
ridges projecting into the nostrils (septal ridges) from the nasal
septum in males; (6) small white tubercles on posterior half of
carapace (males only); and (7) an ovoid carapace (adult and
nearly adult specimens).

Description of holotype. — Carapace ovoid, margin smooth later-
ally, rugose posteriorly; dorsal surface of carapace smooth except
for posterior part; anterior margin of carapace having obtuse
prominences but lacking tubercles; posterior fleshy part of cara-
pace having numerous minute longitudinal corrugations; width of
carapace contained in length, 1.4 times; height contained in width,
2.8 times. Posterior lobe of plastron roimded and truncate; width
of bony bridge, 32 millimeters. Head terminating in elongate
fleshy snout; nostrils rounded, each with a ridge (septal ridge) pro-
jecting laterally from nasal septum; horny parts of jaws concealed
by fleshy lips except anteriorly; internal nares each partly covered
by elongate flap of oral integument projecting from lateral border,
each flap bearing fleshy denticulations on medial edge; least dis-
tance between orbits, 4 millimeters. Forefeet and hind feet fully
webbed, having five digits each; first three digits of each foot bear-
ing claws; four cornified areas (three of which are falciform and
have a free edge) on antebrachium; each hind limb having smooth
cornified area on posterodorsal surface and another (with free
edge) on posteroventral surface. Tip of tail flexible, blunt, pro-
jecting beyond posterior edge of carapace; skin of tail rugous, less so
ventrally than dorsally; anterior tip of cloaca to tip of tail, 20 milli-
meters; posterior edge of carapace to tip of tail, 24 millimeters (see
table 1 for other measurements of holotype and paratypes).

Dorsal surface of carapace blackish in general aspect (dense
mottling of blackish brown and gray evident when specimen is im-
mersed in preservative), lacking pale marginal band; plastron and
undersurfaces of carapace whitish, having numerous small blackish
marks; blackish marks more numerous in area of bridge than on
other ventral surfaces; hyoplastra, hyjooplastra, and xiphyplastra
(plastral callosities) bluish gray where visible beneath translucent
skin of plastron; snout and side of head bluish gray; head and neck
blackish above, bluish white with fine scattered darker markings
below; no pattern on snout or side of head; limbs slate with paler

A New Softshell Turtle (Genus Trionyx) 23

areas of bluish gray above, whitish and tinged with red below;
ventral surfaces of hind feet speckled with black; inguinal area
whitish, tinged with red, lacking darker markings; tail slate above,
whitish and tinged with red below.

Variation. — The paratypic series consists of one adult male (ma-
ture, as indicated by elongate preanal region with cloaca extending
beyond posterior edge of carapace, and fully developed penis ) and
eight subadult females.

The male (PI. II) is unique in lacking septal ridges. The cara-
pace is dark gray and has small white tubercles posteriorly. The
anterior edge of the carapace is smooth. The posterior edge of the
carapace has an obscure pale narrow band, and lacks the corruga-
tions seen in females. The over-all dorsal coloration is blackish.
The ventral surface is whitish with a few black marks on the under-
side of the posterior flap of the carapace. The ventral surfaces of
the tail and hind limbs are tinged with red.

The seven females resemble the holotype in over-all blackish
dorsal coloration, obtuse prominences on the anterior edge of the
carapace, and lack of a pale band on the posterior edge of the
carapace. Shape of carapace varies from ovoid (KU 46908) to
nearly circular (KU 46909). Four of the specimens (KU 46904,
46908, 46910, and 46912) difter from the holotype in lacking longi-
tudinal corrugations on the posterior part of the carapace. On the
smallest female ( KU 46904 ) a dark line extends anteriorly from each
eye and a dark line connects the anterior margins of the orbits.
Black pigment on the ventral surfaces is reduced in three speci-
mens (KU 46904, 46910, and 46912). The ventral surface of the
tail is especially reddish in two specimens (KU 46910 and 46912).
Flaps of skin extending from the lateral borders of the internal
nares partly cover the nares of all KU paratypes as well as the holo-
type. Flaps occur also in the specimens of T. spiiiifcr cmoriji ex-
amined by us (as well as in other forms of Trionyx examined by
Webb); the flaps in cmoriji differ from those of ater chiefly in being
folded vertically against the lateral borders of the nares (not ex-
tending horizontally into and partly covering the nares). Possibly
the flaps are movable; as yet, we are unable to evaluate the func-
tional and taxonomic significance of these flaps.

One female paratype (lU 43510) more closely resembles T. spini-
fer emoriji than do other specimens of T. ater. Pertinent features
of the paratype are as follows: (1) more prominent mottling of
carapace and other dorsal surfaces; (2) posterior rim of carapace
having obscure, pale, narrow band; (3) snout having obscure.

24 The Unin'ersity Science Bulletin

emoryi-hke pattern ot dark lines; (4) plastron less extensively
blackish; and (5) internal narial flaps vertical, not projecting into
internal narial openings. Possibly the specimen is a hybrid.

Range. — Known only from the tyjje locality and other ponds in
the basin of Cuatro Cienegas, central Coahnila, Mexico. Miiller
(1878:641) listed Anujda miitica from "Mexico" but did not men-
tion a specific locality. Presumably Miiller referred to a male of
T. (iter (T. iiiutictis is not known to occur in Mexico). The present
report brings to two (T. ater and T. spinifer emoriji) the numlier of
kinds of softshell turtles known to occur in Mexico.

Relationships. — Trionyx ater can be distinguished from all odier
American forms of the genus by the following combination of
characters: (1) an over-all blackish, dorsal coloration; (2) lack of
all but a trace of a pale marginal rim on the carapace; (3) the lack
of septal ridges in males; and (4) the presence of longitudinal cor-
rugations on the posterior part of the carapace. We consider the
closest living relative of T. ater to be T. spinifer emoriji. Both
forms resemble T. muticus by virtue of reduction in size (emoryi)
or complete loss (males of ater) of tubercles on the anterior edge
of the carapace. Loss of the septal ridge in males of T. ater tends
also to ally this species with T. muticus (septal ridges lacking in
both sexes).

Trionyx ater is closely related also to T. ferox of Florida. Both
species have an over-all slate or blackish dorsal coloration, lack a
well-defined pattern on the limbs, and have at most a narrow or
obscure pale marginal band on the carapace. In both species the
carapace is ovoid and never has tubercles that are sharp-pointed
or conical on its anterior edge. T. ater differs from T. ferox in ha\'-
ing longitudinal corrugations on the posterior part of the carapace
and no septal ridges in males. The corrugations suggest but differ
from the longitudinal rows of tubercles in ferox. Both species have
relatively restricted, southerly displaced, geographic ranges.

It is of interest that the male of T. ater, having a smooth anterior
edge of the carapace, and no ridge projecting from the nasal septum,
resembles T. muticus more closely than do the females of T. ater.
Females of T. ater have a suggestion of tubercles along the an-
terior edge of the carapace, no pale marginal rim on the carapace,
and corrugations on the periphery at the posterior end of the cara-
pace, thus resembling T. ferox more closely than does the male.

Trionyx ater seems to be a relict population of pre-spinifer stock.
Although the resemblance and relationship of T. ater and T. s.

A New Softshell Turtle (Genus Trionyx) 25

emonji is close, we consider atcr a full species because of its geo-
graphical sympatry with T. s. emoriji and the apparent lack of popu-
lations intermediate between the two forms.

The sympatry of T. s. emonji and T. atcr is accompanied by par-
tial ecological separation. Habitat preferences of the two species of
Trionyx in the basin of Cuatro Cienegas were evident in that only
one specimen (KU 46907, adult male, PL II) of T. spinifer emonji
was obtained in the non-fluviatile water of the type locality of
T. (iter, whereas specimens of both sexes of emonji [KU 46913-16
(and two specimens that escaped) PI. I] but no specimens of T. a'^er
were obtained in the Rio Chicjuito, 10 kilometers south of Cuatro
Cienegas. At the latter locality the river is 30 to 50 feet wide and
has a swift current; \ egetation of the quiet backwaters is much like
that of the type locality of T. ater however ( see discussion of habits
and haljitat). Much the same habitat relationship exists between
T. ferox and T. s. asper in areas where the ranges of the two species
overlap. Crenshaw and Hopkins (1956:16) stated, regarding the
area of overlap of these two forms, ". . . asper is nearly always
an inhabitant of fluviatile situations whereas ferox is equally closely
confined to non-fluviatile lakes and ponds."

Studies of other species of acjuatic vertebrates from the basin
of Cuatro Cienegas indicate that the pond habitats in the basin wer(>
isolated in the past and that because of this isolation speciation
of vertebrates took place to varying degrees. The present external
drainage of the area has permitted o\erlap and subsequent inter-
breeding of some of the previously isolated forms with those from
the Rio Salado and Rio Grande drainages, whereas certain other
species, through ecological specialization have been able to remain
distinct. T. ater seems to be in the latter category. The fact that
ecological separation of T. atcr and T. s. emonji is not complete in-
dicates that interspecific matings, possibly resulting in hybrids,
might occur. Matings of T. spinifer wilh T. muticus have been re-
ported (Legler. 1955).

One of us ( \Wbb ) is currently undertaking a taxonomic study
of American forms of Trionyx.

Habitat and habits. — Cuatro Cienegas is situated in an intermon-
tane basin, the floor of which has an ele\'ation of approximately
2400 feet. The basin is approximately 30 miles long (from west
to east) and five to 15 miles wide. The Rio Chiquito (called "Rio
Colorado" by some natixes), originates in the southwestern part,
receives several intermittent tributaries within the basin, and Hows
out through a gap in the eastern end. The Rio Chiquito flows

26 The University Science Bulletin

thence northeastward and joins the Rio Salado near Hermanas,
ultimately draining into the lower Rio Grande near Zapata, Texas.

Much of the central part of the basin is marshy. The sandy
slopes that lead up to the rocky sides of the valley are dry. A num-
ber of clear, deep ponds of various sizes, chiefly west and south of
the town — as well as a hot spring approximately nine kilometers
south of town — occur in the marshy areas. The ponds were pre-
viously isolated but are now drained ( and interconnected ) by small,
man-made ditches that lead to larger cement-lined ditches. It was
learned from natives that water remains in the ponds and in the
Rio Chiquito at all times of the year. Gilmore (1947:148-150,
fig. 2) presented a brief but adequate general description of the
valley.

The type locality of T. (iter is a pond, having a surface area of
approximately two and one-half acres, in the east-central part of
the valley. The average depth of the pond was estimated to be
six feet; holes two to three times as deep were observed in several
places. Water in the pond is warm (approximately 80 degrees
Fahrenheit ) and clear, enabling one to see the bottom clearly in the
deepest places. A narrow intermittent channel leads from the
northern side of the pond toward the Rio Chiquito, approximately
four miles distant. Thick patches of submergent aquatic vegeta-
tion (chiefly stonewort, Chara zelandica var. incoiistans) cover ap-
proximately half of the bottom; the remaining areas are bare, re-
vealing a grayish sediment. Water lilies (Nymphaea sp. ) grow in
the shallower parts of the pond and thick stands of cattails (Typha
latifoUa) and spike-rushes (Eleocharis rosiellota) grow in and near
the water at the edges of the pond. The nearly flat area surround-
ing the pond is grassy or marshy for a distance of several hundred
yards and then gives way gradually to xeric associations including
cacti, yucca, and mesquite. The I. U. paratv'j^e was obtained in a
similar but much smaller pond on the northern side of the basin.

Specimens of T. ater were captured, along with large numbers of
Pseudemys scripta, in hoop-nets baited with canned sardines, rodent
bodies, and fresh cichlid fish. Some turtles of both species entered
the hoop-nets as soon as 30 minutes after fresh bait had been placed
in the nets. Examples of T. ater seemed to be attracted bv the small
live fish that entered the nets as well as by the bait. The relative
abundance of turtles in the pond, reckoned on the basis of trapping
records, may be expressed as one individual of Trionyx per 4.2 in-
dividuals of Pseudemys.

A New Softshell Turtle (Genus Trionyx) 27

The heads of several T. ater could usually be seen at dusk by
scanning the surface of the pond with binoculars. No turtles were
seen on the surface or beneath the water during daylight hours;
it is assumed they sought cover beneath dense vegetation or in sedi-
ment at the bottom of the pond. Softshells entered hoop-nets at
^'arious times between dusk and dawn and continued to enter in the
course of the day if the sky was overcast; success of trapping de-
creased sharply during periods of bright sunlight.

The stomach of one of the paratypes (KU 46908) contained 23
larvae of long-eared leaf beetles (Chrysomelidae: Donacia sp.)
and many short pieces of roots of spike-rushes; the roots probably
were ingested incidentally along with the larvae that were attached
to them.

Remarks. — The second and third largest paratypes (KU 46906
and 46908) were dissected and found to have immature ovaries;
probably, therefore, the holotype, a slightly larger specimen, is
immature as well. Larger adults were probably unable to enter
hoop-nets, the largest of which had openings slightly less than one
foot wide.

Natives at Cuatro Cicnegas refer to softshell turtles as "tortuga
hlanca"; they make no distinction between T. spinifer etnoryi and
T. ater.

ACKNOWLEDGMENTS

We are grateful to Dr. Hobart \L Smith and Messrs. Pete S.
Ghrapliwy and Kenneth L. Williams for permitting study of ma-
terial at the University of Illinois; to Messrs. Wendell L. Minckley
and Robert B. Wimmer for assisting Legler with field work; and to
Drs. George W. Byers and Ronald L. McGregor, respectively, for
identification of insect larvae and plants. Special thanks are due
Sr. Daniel Rodriquez Villarreal, Cuatro Cienegas, who gave gen-
erously of his time in guiding the K. U. field party to the type lo-
cality.— Museum of Natural History, University of Kansas.

LITERATURE CITED

Crenshaw, John W., and M. N. Hopkins, Jr.

1955. The relationships of the soft-shelled turtles Triontjx ferox ferox
and Triomjx ferox uspera. Copeia, 1955, No. 1, pp. 13-23, 4 figs.,
February 18.
GiLMORE, Raymond M.

1947. Report on a colleetion of niainnial bones from archeologic cave-
sites in Coahuila, Mexieo. Journ. Manun., vol. 28, No. 2, pp.
147-165, 1 pi., 2 figs., 1 table. May 19.

The Univfrsh V Science Bueletin

Legler, John M.

1955. Observations on the sexual behavior ot captive turtles. Lloydia,
vol. 18, No. 2, pp. 95-99.

Mi'LLER, R.

1878. Katalog der im Museum unci Universitiitskabinct zu Basel aufgestell-
ten Amphibien und Reptilien nebst Annierkungen. Verb. Naturf.
Ges. Basel, vol. 6, pp. 557-709, 3 plates.

Measurements (in millimeters) ot type and paralypes ot Triontjx ater new
.species. Length of snout was measured from middle of tip of snout to an-
terior corner of eye. All measurements are maximal.

Length

Width

Length

Width

Length

Number

Sex

Height

carapace

plastron

head

snout

KU 46903

243

178

183

KU 46906

231

181

172

KU 46908

213

148

155

KU 46910

211

152

158

lU 43510

210

164

163

KU 46905

205

156

151

KU 46912

189

140

139

KU 46909

186

142

136

KU 46904

152

117

108

KU 46911

134

108

A New Softshell Turtle (Genus Trioxyx) 29

PLATE I

Top. Trionijx ater sp. nov., dorsal and ventral views of holotype, KU 46903
(X Vb). Bottom. Trionyx .spinifcr cnionji, dorsal and ventral views of female
(KU 46913) from Rio Chiquito, 10 km. S Cuatro Cienegas, Coahuila, Mexico

The University Science Bulletin

PLATE II

Top. Triontpc ater sp. nov., dorsal and ventral views of adult male, KU
46911 (X y*)- Bottom. Triomjx spinifer emonji, dorsal and ventral views of
adult male (KU 46907) from type locality of T. ater (X^)-

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 3

A New Caecilian Genus in India

Edward H. Taylor

The Caecilian fauna of Pakistan and India has long been known
to comprise species of four recognized genera: Ichthyophis Fitz-
inger, UraeotypJihis Peters, Gegeneophis Peters, and Herpele Peters.
A fifth genus is herein described. Some of the more obvious dif-
ferences in the five genera may be expressed in the following key:

Key to Asiatic Caecilian Genera

1. Scales absent; one row of teeth in the lower jaw; tentacle cone-shaped,
close behind and below nostril, nearer to tip of snout than to eye;

squamosal and parietal bones in contact Gegeneophis

Scales present; one or two rows of teeth in the lower jaw 2

2. Eye hidden, covered by bone; tentacle conical, below and somewhat
behind the nostril, closer to tip of snout than to eye, the tentacular

opening circular; squamosal and parietal bones in contact Herpele

Eye not covered by bone, more or less visible; tentacle \'ariable 3

3. Vent transverse; tentacular opening horseshoe-shaped, surrounded by a
low craterlike elevation whose inner sides are marked with minute
valleys and ridges ( shape of extruded tentacle not known ) ; tentacle
situated directly in front of eye and on a level with nostril, but much
closer to eye than to nostril, the tentacular groove on skull forming a
forward extension of the orbit; (relation of skull bones to each other

not known ) ; no tail present, end of body rounded Ii}dott/})]iIus

Vent longitudinal, a short pointed tail,, bearing transverse folds 4

4. Tentacle flaplike, nearer to tip of snout than to eye, situated almost di-
rectly below nostril, near mouth; squamosal and parietal bones sepa-
rated by a diastema; orbit not elongated forward; a prefrontal and
postfrontal; no frontoparietal foramen; internal nares open between

vomer and palatine; fewer than 200 folds, none anguhu Uraeotyplilus

Scales present under skin throughout body; tentacle conical, near edge

of mouth, much closer to eye than to nostril; parietal and squamosal
bones in contact; more than 200 grooves (or folds); one or two rows
of teeth on mandible; folds on venter largely angular Ichthyophis*

* There is strong probability that when it is possible to study skulls of the various
species now described, Ichthyophis will he found to consist of more than a single genus.

(31)

The University Science Bulletin

Fig. 1. Indotyphlu.s battcrsbyi sp. nov. Photograpli oF tlic type specimen.
Actual length, 170 mm. Khandala, Foona District, India.

A New Caecilian Genus in India 33

Indotyphlus^' gen. nov.

Type of genus: IndotypJihis battershyi sp. nov.

Diagnosis: Anal vent transverse; no tail, the postanal region
blunt, rounding, lacking transverse folds; 139 primary folds sur-
round body; secondary folds present in posterior part of body only;
eye concealed below skin and flesh; the orbital rim incomplete, the
tentacular depression of skull continuous with the orbit; eyeball
more or less imbedded on the base of tentacle which runs straight
forward from orbit on level with nostril but emerges distinctly closer
to eye than to nostril; vertebrae 144. A maxillary-premaxillary and
vomeropalatine series of teeth in upper jaw; lower jaw with an
outer mandibular series and a shorter inner splenial f series.

Indotyphlus batfersbyi sp. nov.

Type: No. 49974 American Museum Natural History, from Khan-
dala, Poona District, India.

Diagnosis: A slender species, the width in length approximately
46 times; secondary folds begin at about the 110th primary fold
and scales present posterior to this point; eye concealed; tentacle
on level with eye, nearer eye than nostril; elevation about tentacle
craterlike.

Description of type: Body slender elongate; snout rather acu-
minate, projecting .75 mm. beyond tip of lower jaw; eye com-
pletely concealed below skin and flesh; tentacular opening, some-
what horseshoe-shaped, appearing at the bottom of a shallow
crater whose inner walls are sculptured with ridges and valleys;
(shape of exserted tentacle unknown but probably conical); width
of head at mouth angle (3 mm.) slightly less than head length
from mouth angle to tip of snout (3.25 mm.); length to tip of snout
from first (incomplete) nuchal groove, 4.2 mm.; length from second
(complete) groove, 5.6 mm.; from third (incomplete) nuchal groove
limiting pharyngeal region, to tip of snout, 7.25 mm.

A short dorsal groove between first and second transverse nuchal
grooves; a short lateral groove indicated between the second and
third grooves; transverse costal (primary) folds and grooves sur-
round body without forming a ventral angle, distinct except for a
space in median dorsal area of back; primary folds, 139; the sec-
ondaries in posterior part of the body begin as short lateral grooves

* From Indo, referrinE to India, + typhlos Gr. = blind.

\ In caecilians the inner inandilinlar series of teeth seemingly is equivalent to the
sjilenial teeth of salamanders. In this i^aper they are called splenial teeth.

2—8920

The University Science Bulletin

at the 110th fold and continue to back edge of vent growing longer,
totalling about 29, only the last 13 surround the body completely;
vent transverse, the edges denticulate; without grooves or folds;
behind vent, the end of body blunt, rounded, its ventral surface a
little flattened.

Fig. 2. Radiograph of the type of Indotyphlus battersbyi sp. nov.

Snout projecting a little beyond mouth; maxillary-premaxillary
tooth series, 9-9 (allowing for missing teeth), the teeth nearly triple
size of those in vomeropalatine series which number, 12-12; man-
dibular series, 9-9, distinctly larger than maxillaries; splenial
series, 2-2, smaller than mandibular teeth; tongue small, pointed,
not covering the splenial teeth; internal nares large, separated by a
distance less than transverse diameter of one choana.

Glands of skin more or less equally distributed over body; a
lateral ridge present, broken by grooves, usually distinct through-
out; scales present in posterior part of body where secondaries are
present, two series for each primary fold; each series with three

A New Caecilian Genus in India

rows of imbricating scales that overlay each other; posteriorly ex-
tending entirely around the body; vertebrae 144.

Color ( in preservative ) : Brown of a nearly uniform shade on
dorsum and sides; head darker; slightly lighter brown ventrally;
a somewhat lighter area near vent.

Fig. 3. Diagram of the liead and anterior part of the body of Indotyphlus
battersbyi sp. nov., showing the relative position of nostril and tentacle. The
eye is not visible. From type, X 12.

Measurements in mm.: Total length, 170; head width, 3.5; head
length, 4.7; body greatest width (approximately), 3.7; width in
length approximately 46 times.

Remarks: The detailed characteristics of the skull are as yet
unknown. When this can be studied, more complete knowledge
of the relationships of Indotyphlus with other genera will be learned.

The type may be one of the specimens collected by Charles
McCann at Khandala, Poona District, and reported by him as Ich-
thijophis monochrous Boulenger, in the Journ. Bombay Nat. Hist.

The University Science Bulletin

Soc, vol. 31, no. 4, Feb. 20, 1927, p. 1039. He states "When at
Khandala during the month of September, 1919, I secured several
specimens of this batrachian. It lives under stones during the
rains in burrows much after the fashion of the earthworm which it
also resembles in its movements. At first sight it might well be
mistaken for one of these creatures as its body is also coated with
slime. On the removal of the stone under which it lives the animal
soon begins its descent into its burrow away from the light."

Fig. 4. Terminus of the bo(l>- of Indotyphlus battcisbyi sp. nov., showing
absence of a tail and the transverse vent. From type, X 12.

In the same Journal, vol. 42, part 1, 1940, on p. 64 Mr. McCann
writes: "On the 6th September 1931 while collecting frogs I dis-
covered another specimen (of Ichthijophis monochrous) living
under a stone on the banks of the lake behind a range of hills lo-
cally called the 'Sausages.' I have repeatedly hunted for this ani-
mal since its first discovery at Khandala, but without much success.
The 1931 specimen measured 232 mm."

I suspect that this second report represents a totally different
species.

The species is named in honor of Mr. J. C. Battersby of the
British Museum of Natural History.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 4

On the Caecilian Species Ichthyophis glutinosus and

Ichthtjophis inonochroiis, with Description

of Related Species

Edward H. Taylor

Abstract: The Asiatic forms of the gcmis Ichthijopliis are considered. The
species generally recognized as Ichthyophis ghitinosus and Ichthyophis mo-
nochrous were each found to consist of several species. The species described
by Taylor, Ichthyophis glandulosus and Ichthyophis xveheri synonymized by
Inger are resurrected and recognized, and the following species synonymized
by Boulenger and others are revived: Ichthyophis tricolor Annandale, /. bed-
domei Peters. Ichthyophis monochrous Bleeker is recognized and is belie\Td
to be confined to the island of Borneo. 7. ghitinosus Linnaeus is likewise rec-
ognized as an Asiatic species.

Larvae from a number of places were examined. Certain of these seemingly
belong to species whose adults have not been seen and are probably un-
described. No attempt has been made to utilize larvae as types of species.

The following fomis are described as new species: Ichthyophis dulitensis,
I. lanitensis, I. paucidentuhis, I. singaporensis, I. peninsularis, I. subterrestris,
I. bomhayensis, I. mindanaoensis, I. javanicus, I. mcdabarensis, I. youngorum,
I. sikkimcnsis, I. sumatrciniis, I. acumiruitus, I. nigrofiavus, I. paiicistdcus, I.
sttpacliaii, I. kolitaocnsis.

A number of specific characters have been considered that have been
heretofore largely neglected. These are the character and distribution of the
scales, the continuity or discontinuity of th& folds and grooves around tlie body,
the presence or absence of a row of splenial teeth (second mandibular series),
the number, relative size and characteristics of the other dental series, and the
length and eharacteristics of the tail.

In tlie lar\ae the lateral line system (neuromast) \aries much and seems
to reflect habits of the young. Certain of the lar\ae ha\e onl>- a single gill
s'it; others have two well-developed slits.

(37)

38 The University Science Bulletin

TABLE OF CONTENTS

PAGE

Abstract 37

Introduction 38

Acknowledgments 43

Taxonomic Consideration; Species of Ichthyophis

I. weberi Taylor 43

I. larutensis sp. nov 44

I. species ( larvae no. 1 ) 47

I. species ( larvae no. 2 ) 49

I. paueidentulus sp. nov 49

I. monochrous ( Bleeker ) 51

I. singaporensis sp. nov 56

I. dulitensis sp. nov 58

I. peninsularis sp. nov 61

I. subterrestris sp. nov 65

I. bombayensis sp. nov U7

I. niindanaoensis sp. nov 69

I. glandulosus Taylor 74

I. javanicus sp. nov 77

I. inalabarensis sp. nov 80

I. youngoruni sp. nov 84

I. sikkimensis sp. nov 91

I. sumatranus sp. nov 95

I. acuminatus sp. nov 98

I. nigroHavus sp. nov 101

I. paucisulcus sp. nov 103

I. supachaii sp. nov 107

I. kohtaoensis sp. no/ 110

I. beddoniii Peters 113

I. tricolor Annandale 113

Bibliography 114

INTRODUCTION

In the work of Seba^ (Thesaurus, vol. 2, 1735) a species of
caecihan was described briefly under the name Serpens caecilia
ceylonica, Ceylon being the type locality of the species. Here
it was presumably considered as a snake.

Nineteen years later, in 1754, Linnaeus listed two caecilians in a
work, "Museum S. R. M. Adolphi Friderici." . . . These were
Caecilia tentacidata from South America and Caecilia ^Jutinosus,
a new species described with "Habitat in Indiis."

The description of glutinosa, published both in Latin and Swedish,

I. Seba, A. Locupletissimi naturaliuin tht-sauri accurata descriptio et iconibiis arti-
ficio sissimus expressio, per universam physices historiani. Vols. 1-4, 1734-1763, 449 pts.
Amsterdam. (Vol. 2 published in 1735.) A copy of this description, vol. 2, p. 26, was
published in Taylor, Univ. Kansas Sci. Bulletin, vol. 38, pt. 2, no. 13. Mar. 20, 1958.

On the Caecilian and Related Species 39

though brief, is scarcely sufficient to distinguish an Indian or Asiatic
species. The description in Latin reads as follows:

Caecilia glutinosa

"glutinosa Caecilia rugis transversalibus 340, caudalibus 10.

"Caput parvum, laeve. Narium foramina in antica parte capitis. Cirrhos
nuUos observare potui. Oculi minutissimi, mcmbrana abdvicti. Denies utrius-
que maxillae minimi, serie dnplici.

"Tnincus cylindricus, \ersus posteriora paiilo crassior: Rugis minutissimis,
vix visum subeuntibus, a tergo retrorsum nonnihil flexis at futuram abdominis,
coeuntes angulo acuto. Rugae majusculae circiter 350.

"Cauda brevissima acutiuscula. anus sub & juxta apicem caudae.

"Color fuscus: linea utrinque laterali albida, latiuscula.

"Longitudo pedem superat. Crassities digiti minimi.

"Glutine vedetur viva esse obducta uti petromyzon."

Further data from the type specimen is necessary before this
species can be placed properly.

Anderson (1899), Catalogue of Linnean type specimens of
Snakes, Bihang Till K. Svenska Vet.-Akad Handlingar, Band 24; Afd.
4, No. 6, page 6, states that 355 folds are present and that the
length is 400 mm.

The description is scarcely adequate for distinguishing a species
of Indian caecilian. Linnaeus states that he has not observed a
tentacle on the head; however one must presume that a tentacle is-
present. That it is not a larva is presumed by the fact that neither
gill slits ( spiracles ) nor a tail fin are mentioned. The species under
the same name is listed by Linnaeus in tlie 10th edition of the
Systema Naturae (p. 229).

Certain subsecjuent authors ha\e used other names presumably
in reference to the Linnaean species. Thus Latreille lists Caecilia
viscosa as a substitute name - and Cuvier uses the name PCoecilia
bivitatta in the Regne Animal (1817, p. 100) while on another
page (tome 2, p. 87) the Linnaean name is used.''

In 1826 Fitzinger ^ proposed the taxon Nuda for his fourth tribe
(Tribus) of the Reptilian, Order (Ordo) Menopnoa. The first and
only family designated in this was Coecilionideo. He included in it
the genera Caecilia Linnaeus [= Coecilia Cuvier] (using the spell-
ing given by Cuvier rather than that given by Linnaeus), and a new
genus Ichthyophis. Of these he states:

2. Latreille, P. A. In Sonini and Latreille, Histoire naturelle des Reptiles. Vols. 1-4.
1802. p. 236.

3. Le regne animal distribue d'apres son organization, pour scrvir de base a I'Histoire
Naturelle des Animaux et d'introduction a I'Anatomie comparee, vols. 1-4, 1817, Paris.

4. Neue Classification der Reptilien nach ihren natiirlichen verwandschaften nebst einer
verwandschafts — tafel und linein verzeichnisse der Reptilien — Sammung des K. K. zoologischen
Museum's zu Wien". 1826, pp. 1-66, 1 table. F. G. H( ubner, Wien.

40 The University Science Bulletin

Isolirt steht diese Familie in der Zunft der Nuden, das Bindungsglied der
Monopnoen mit der Dipnoen bildend. Beim Ueberblicke der Arten der
Cuvier'schen Gattung Coecilia zeigt sith sogleich in der Totalform cine Ilaupt-
verschiedenheit, welchc zur Trennung meiner Gattung Ichthyophis Veran-
lassung gegeben hat. Linne's Caecilia glutinosa und eine Art aus Java haben
einen deutlich plattgedriicktcn Korper mit zugespitztem Ende, wcleher den
iibrigen Coecilen durehaus fehlt, und hierauf ist der generisliee Unterschied
gegrijndet. Coecilia bildet den schonsten Uebergang zu Leposternon aus der
Zunft der Squamaten, nnd durch Ichthyophis zu Amphiuma, Siren und Pseudo-
branchus, aus der Zunft der Imniutabilien in der Ordnung der Dipnoen."

Subsequent to the proposal of the genus IchthyopJtis; Fitzingor
proposed the name Ichihyophis Hasseltii for a species based on a
specimen in the Vienna Museum from Java. One is forced to con-
clude that the Ichthyophis HasseUii of Fitzinger is a nomen nuduDi
since he offers only the following meager data on the specimen :

Page 36: Caecilia glutinosa and a new species from Java have a distinct
"plattgedruckten" body v^ath a "zugespitztem Ende."

Page 63. Fitzinger hsts species present in the Vienna Museum: Coecilia
anntdata and C. lumhricoides respectively from Brasil, and an imknown lo-
cality; and a third species, Ichthyophis Hasseltii, from Asia, Island of Java.

These data seemingly cannot distinguish hasseltii from certain other species
of Ichthyophis now known.

The following year Van Hasselt, in Isis, 1827, p. 565, under the

name Coecilia hijpocijanea described a species from Java in the

Vienna Museum, presumably the same specimen that served as the

basis for the name I. hasseltii. The description follows:

"Capita indistincto, depresso laevissimo, tentaculato, tenlaculo ad marginem
maxillarum utrinque anteorbitali minuto, oculis par\is hebetibus, cauda bre-
vissima, trunco fusiformi, capite paulo latiori, nigis 320 circiter sutura ab-
dominali oblique interruptis, arctissime annalato. Supra ex olixaceo obscura,
subtus chalybea linea laterali flavopunctata;"

The following authors have used this name and figured the form:
Miiller 1835, p. 391, pi. 8, figs. 12-14. Schlegel, 1832-1844, p. 119,
pi. 39, fig. 1. Wagler, 1828, p. 743 reports this species as Epicrium
Hasseltii. "Corpore supra obscure olivaceo, subtus chalybeo-cae-
ruleo, lateribus lineis duabus maculosis flavo ochraceis picto; trunci
rugis circiter 320. Habitat in Ja\a insula locis paludosis Javanis
Octur-doeel dictum."

The original description of Ichthyophis monochrous was pub-
lished by Bleeker (Nat. Tydschr. Nederl. Indie, vol. 16, 1858) as
Epicriwii monochroum, in section 7 of the generalized title —
"Bestuursvergadering gehouden ten huize van den Heer De Bruijn
Kips den 11 Maart 1858."

The section 7 begins: "De Heer Bleeker deelt made, dat hijdezer

On the Caecilian and Related Species 41

dagen weder eene verzameling reptilien en \ isschen van Sinkawang
(westkust van Borneo) ontvangen heeft.

"Deze verzameling is, evenals vroegere van Sinkawang, aange-
boden door het lid der Vereeniging, den heer J. H.A.B. Sonnemann
Rebentisch.

"De reptilien bestaan uit de folgende soorten." . . . [On
page 188 the following description appears]:

"No. 15 Epicrium monochroum Blkr.

"De heer Bleeker vertoont laatstgenoemde soort in een voorwerp
van 232 millimeters lengte. Het is van eene violetbruine kleur over
het geheele ligchaam en slechts iets lichter gekleurd aan de onder-
vlakte van den kop, terwijl de anus door eene witachtiggele vlek
omringd is. De overlangsche gele band van Epicrium glutinosum
Wagl. ontbreekt volkomen. Ook zijn er niet, zooals de heer A.
Dumeril van Epicrium glutinosum opgeeft, 325 volkomene cirkel-
vormige huidplooijen maar slechts 246 plooijen in het geheel, welke
alle aan de buikvlakte een' stompen naar achteren gerigten hoek
vormen. De hoogte des ligchaams gaat ovgeveer 25 malen in zijne
lengte. Het kuiltje onder de oogen bestaat doch is zeer opper-
vlakkig, heeft weinig verhevene randed en ligt nabij de bovenlip
een weinig voor het oog. De schubjes zijn uiterst klein. Het
ligchaam is aanmerkelijk dikker in het midden dan aan het kop-en
staarteinde. De heer Bleeker beschouwt het voorwerp op grond
der genoemde kenmerken als eene eigene soort, welke vrij aan-
merkelijk van Epicrium glutinosum verschilt."

Peters,'' in 1879 described a species from India under the name
Ichthyophis hcddomii while a second Indian form was named Ich-
thyophis iijutinostis tricolor by Annandale in 1909.^

Boulenger in the "Catalogue of the Batrachia Gradientia s Cau-
data and Batrachia Apoda in the collection of the British Museum"
(1882) recognizes only two species in the genus Ichthyophis and
Boulenger has been largely followed by all subsequent writers
dealing with the genus.

Thus the genus Ichthyophis for more than half a century was
generally regarded as comprising only two species of caecilians.
One, the species Ichthyophis ghitinostts, described by Linnaeus in
1758 whose range came to be defined as extending from the East
Indies through southeastern Asia to India. The second species,
Ichthyophis monochrous described by Bleeker in 1858 was sup-

5. Mon. Akad. Berlin 1879, p. 931.

6. Rec. Indian Mus.. vol. 3, 1909, p. 286.

42 The University Science Bulletin

posed to range from Borneo through Ja\a, Sumatra, Malaya, Siam,
Ceylon, and India, to Sikkim.

In 1920 I obtained a specimen of a caecilian discovered by C. M.
Weber ' on Palav^an Island, Philippine Islands. This I described
as a nev\^ species, Ichthyophls tveberi differing, among other charac-
ters, from 7. monochroiis in lacking the inner row of mandibular
teeth (splenials) and in having some 78 more circular folds about
the body *" characters which an experienced taxonomist probably
would accept as being worthy of designating a different species.

In 1923 I discovered still another form of caecilian on Basilan
Island, Philippine Islands, which was named IclitJiyophis glandu-
losus.^

The published ranges of these two species, 7. g,lutinosus and
monochrous, were truly surprising. It became apparent that a re-
markable happening had taken place in the southeastern part of
the range of the genus in the Philippines and Borneo where no less
than three species were present; while no change was recognized
in the two species which purported to extend from Borneo to Sikkim
in the Himalayas, and to Ceylon, a distance of several thousand
miles.

It is almost an axiomatic rule that genera of low vagility tend,
as they spread, to speciate to a greater degree than those of greater
vagility. Burrowing genera of reptiles and amphibians are per-
haps the least vagile of all the vertebrates. But here were two
recognized species of burrowing caecilians whose ranges were
enormous and in which, presumably, there had been little species
formation, thus contradicting the generally accepted belief. This
problem definitely merited investigation, to account, in some way,
for this lack of plasticity in the genus, or to prove that such ranges
did not exist.

On the basis of experience in the Philippines, I concluded that
most evidence pointed to the fact that previous authors had in
some manner confused species (since extraordinary variation in
the number of the primary and secondary folds had been recorded
in the literature), and that we were dealing with species — perhaps

7. Later killed by natives on the nearby island of Balabac.

8. This form has recently been placed by Dr. Inger in the synonymy of Ichthyophis
vjonochrous. This should be disregarded, largely because the author has a different con-
cept of species (his own statement), because of his inexperience with Amphibia, and per-
haps his lack of knowledge that much caecilian material in museums is misidentified.

9. This form also was relegated to the synonymy of /. monochrous by the same
author as the preceding. He comments that the lateral fold is not glandular, despite the
fact that on it many hundreds of glandules are present. He states, "Ichthi/ophui glandttlosus
Taylor is probably conspecific with monochrous Bleeker. I am unable to find distinguishing
characters" . . . this statement I believe, is made by Inger without his having seen a
specimen of either monochrous or glandulosus. His action I believe should be disregarded
for the reasons set forth in the preceding footnote.

On the CAEaLiAN and Related Species 43

even generic complexes in the nominal Ichthyophis monochrous and
Ichthyophis glutinostis.

In 1958 I discovered certain imdescribed species of the genus in
Thailand, definitely not to be associated with either monochrous or
ghitinos'us. On my return journey from Thailand to America I
visited museums in Ceylon, Germany, France, and England and
examined the specimens classified imder the two above names. I
found that my tentative conclusion was confirmed — that several
undescribed species were masquerading in museums under the
names Ichthyophis monochrous and 7. glutinosus.

Through the kindness of the staffs of the various museums I was
permitted to record data or borrow specimens for later study. Fur-
ther material was made available by certain American museums.
There follows here a review of most of the a\"ailable material with
the results of my study.

ACKNOWLEDGMENTS

I wish to acknowledge loan of specimens or other assistance from
the following: Mr. P. E. P. Deraniyagala, Dr. Robert Mertens, Dr.
Konrad Klemmer, Dr. Heinz Wermuth, Miss Alice G. C. Grandison,
Mr. J. C. Battersby, Dr. Doris Cochran, Dr. Ernest Williams, Dr.
Charles Bogert, Dr. Robert Inger, Dr. Alan E. Leviton, Dr. George
Myers, Dr. Walter C. Brown, and Dr. and Mrs. Hobart M. Smith!

TAXONOMIC CONSIDERATION

Ichthyophis xcvhcri Taylor

Ichthyophis tveheri Tavlor, Philippine Journ. Sci., vol. 16, no. 3, March 1920,
pp. 227-228 (type locality, Malatgan River, Palawan, P. I., C. M. Weber
coll.); Dept. Agri. Nat. Resour. Bureau Sci. Manila, Publ. 15, December
15, 1921, pp. 26-27 (reprint of preceding type description).

IchtJitjophis monochrous van Kampen (part.), The Amphibia of the Indo-
Australian Archipelago. Leiden, 1923, pp. 3-4, 282 (svnonomv questioned);
Inger (part.) Fieldiana: Zoology, vol. 33, no. 4, July 23, 1954, pp. 207, 209
(imquestioned synonymy).

Type: Formerly, No. B-1, Bureau of Science collection; collected
at Malatgan River, Palawan, January 28, 1909, by C. M. Weber.

Description of type: "Two rows of teeth in upper jaw, the series
forming oval arches, parallel to each other, the inner row extend-
ing much farther back than the outer but not widening; lower jaw
with a single row of teeth, with no evidence of a second row; head
oval, eyes distinct, the distance between them very slightly less
than width of head between eyes; distance between eyes a little
greater than length of snout; tentacle withdrawn, the groove rather
moon-shaped, situated anterior to eye near the edge of upper jaw;

44 The University Science Bulletin

body surrounded by three hundred twenty-four circular folds meet-
ing on belly in an angle, except those on posterior part of body,
which run straight across without an angle; the first three or four
folds on anterior part of body fail to meet; a more or less distinct
groove from tip of lower jaw to some distance in front of anus along
the median ventral line of body.

Color in alcohol: "Above yellowish brown, somewhat darker on
median part of body; below lighter yellowish brown. Under a
microscope the color appears as minute rounded yellowish dots
surrounded by a network of brown. A white spot on tip of lower

jaw."

Measurements of Jchihyophis weberi Taylor

mm.

Total length ^ 250.

Tail 2.5

Width of head at eyes 7.5

Length oi snout 5.

Eye to nostril 3.5

Eye to tentacle 1.5

Remarks: To the best of my knowledge no specimen other than
the type has ever been taken of this species. I know of no scientific
expeditions that have been undertaken in the island of Palawan in
recent years.

I visited the Bureau of Science in Manila in September 1957 with
the intention of re-examining the type specimen of Ichthyophis
weberi, only to learn that the type together with all the extensive
herpetological collections of the Bureau of Science, largely brought
together by me, had been destroyed during the second World War.

Ichthyophis lanitensis * sp. nov.

Ichthyophis monochrous Flower** Proc. Zool. Soc. London, 1899, (Nov. 14),
p. 916 (not ol Bleeker).

Type: British Museum Natural History, No. 98.9.22.208, Larut
Hills (near Maxwell's Bungalow), Perak, Malaya, elevation 3,380
ft. Stanley Smyth Flower collector. (A second specimen men-
tioned, see footnote. )

* Larut ( Mts. ) + ensis ( Latin ) = place, locality, country.

** Flower comments on the form: "In April 1898 on different clays I obtained two
specimens from under a stack of firewood near 'Maxwell's Bungalow' in the Larut Hills,
Perak, elevation 3380 feet.

"1st. Number of circular folds about 313; length 208 mm.

"2nd. Number of circular folds about 309; length 167 mm.

"As in I. glutinosus, some of the circular folds either bifurcate or converge into each
other; therefore the number, in counting the same individual at different parts of its cir-
cumference, varies.

"Colour (in life): Uniform purplish black. Tentacles white. Anal region and tip of
tail pale pinkish. The eye appears as an inconspicuous black speck (but turns whitish in
spirits)."

On the Caecilian and Related Species

Fig. 1. IchthyopJm larutensis sp. nov. Type B. M. N. H. No. 98.9.22.208
Larut Hills, Perak, Malaya, elev. 3380 ft. Actual length, 169 mm.

46 The Uni\'ersity Science Bulletin

Diafftiosis: A slender species, the body width in length. 30.7
times; tail length in body length, 48.3 times; transverse folds do not
cross back in anterior three fourths of the body; approximately 304
transverse folds ("about 309" counted by Flower loc. cii.)\ folds
on tail, 7; no splenial teeth present; tentacle nearer to eye than to
nostril. Vertebrae, 107.

Description of type: Head slender, oval, the width at first nuclial
groove, 5.1 mm.; the head length, 11 mm.; eyes distinct, the
pupil white, surrounded by a narrow black iris; this in turn sur-
rounded by a narrow cream ring; distance between eyes, measured
on the curve, 3.8 mm., a little broader than length of snout (3.5
mm. ) ; tentacle close to lip, nearer to eye ( 1 mm. ) than to nostril
(1.8 mm.).

First transverse nuchal groove crosses throat, runs upward and
somewhat posteriorly, but fails to cross the head completely; the
second groove crosses throat and goes up on side of neck nearly to
dorsal level of neck; the third groove is distinct ventrally only and is
rather widely separated from the second, forming a median angle
below; primary and secondary grooves and folds about 304; they
do not cross the middle line of the back except on the posterior
fifth of the body; medially they fail to cross venter completely on
anterior part of body; eight folds on tail counting from front of
vent; tip of tail rather sharply pointed; a slight longitudinal groove
on back part of chin connecting with the second groove.

Scales are absent, or minute and scattered in the anterior third
of the body; posteriorly they are larger but consist usually of a
single overlapping row in each fold.

Teeth: maxillary-premaxillary, 21-20, those on the premaxillae
largest; vomeropalatine, 19-19; mandibular, 19-19; splenial, 0-0;
tongue well developed covering entire area between maxillary-pre-
maxillary series, not suggesting a recent transformation from the
larval state; vertebrae present, 107.

Color in life (fide Flower): "Uniform purplish black, tentacles
white; anal region and tip of tail pinkish;" at present the general
color is dark brown; a cream-white spot on tentacular area and
one about nostril; the grooves on throat are cream; lips lighter than
head; the tip of tail and anal region cream.

Measurements in mm.: Total length, 169 (fide Flower, 167); tail,
3.5; body width, 5.5; width of head, 5.1; length of head, 11.

Remarks: The failure of the transverse folds and grooves to cross
the back except in the posterior fourth is a significant character.

On the Caecilian and Related Species

The light marks on the throat, the character of the third groove,
the absence of the splenial teeth, and the pink spots at vent and
tail-tip clearly delineate this form.

Two larval species of caecilians obtained in Perak are treated
here. I am convinced that neither are the young of Ichthyophis
larutensis.

Fig. 2. Ichthyophis larutensis sp. nov. Type. Radiograph
showing 107 vertebrae

Ichthyopliis si^i. (1)

A series of fifteen larval specimens taken "near Yum River, head-
waters Plus River, East Perak, Malaya, 2000 ft." and bearing the
British Museum numbers 1935.5.21.51 to 1935.5.21.65, cannot be

The Uni\'ersity Science Bulletin

placed definitely in a recognized species. In fact it wonld appear
that two forms are represented in the material, and neither of them
is the yonng of the species described here as larutemis.

The specimens were all taken in March 1923. They resolve
themselves on the basis of length measnrements into possibly three
(presumably age) groups as indicated in the following table. These
groups are not clearly limited, a fact which suggests that the re-
production period during a year is a rather extended one.

The table includes data on a second larval form (B. M. 1934
.5.21.50) and the type of Ichthyophis larutensis.

Data on two larval forms of Ichthyophis, and I. larutensis sp. nov.

Number

bfl

a
-a

a
u

Ma.x-premaxil-
lary teeth

t4
Is

"a
m

1934.5.21.65

56
62
65
65
70

1934.5.21 60

98
103

4.8
5

20
20.5

244
255

12-14

15-16

13-13

4-4

113

5.7

247

13-14

15-15

13-13

4-4

115

240

13-14

17-17

13-14

4-4

122

266

15-14

15-17

15-16

4-5

1934.5.21.56

132

249

13-13

18-18

16-15

5-5

137

7.3

18.7

257

15-15

21-21

17-16

5-5

140

7.2

261

14-14

18-18

15-15

5-5

140

7.1

268

15-14

18-18

5-5

141

7.2

265

13-14

17-18

14-15

5-5

1934.5 21.50*

184

348

13-14

18-18

20-20

1-1

98.9.22.208**

169

5.5

304

21-21

20-19

20-20

0-0

* Second larva.

** Type of larutensis.

The folds and the teeth on the youngest specimens cannot be
counted. The eyes are indicated by rounded milky-colored areas,
through which the eyeball is rarely visible. The tentacular opening
is indicated either by a shallow pit or an opening (in older speci-
mens) in the anterior part of eye-spot. The tail fin begins on the

On the Caecilian and Related Species 49

dorsal part of the body about one or two millimeters in advance of
the vent and continues around the end of the tail then forward to
the back end of vent. The nostrils are terminal and not visible di-
rectly from above.

The gill opening is single with short free flaps preceding and
following it; the transverse folds meet in the median ventral line
at a sharp angle throughout most of the body except the part in
front of the vent where they are almost straight across the venter.
Six or seven folds are on the tail (counting from the front of the
vent). The variation in the number of body folds in the specimens
counted vary from 240 to 268. It is belie\ed that much of this
variation is sexual, females having the higher counts in at least
some species.

Ichthyoplns sp. (2)

A single specimen, B. M. No. 1934.5.21.50, taken in the locality
"near Yum River, headwaters of Plus River, East Perak, 2000 ft."
Malaya, differs from the preceding series in such a way as to sug-
gest that it represents a different species.

The chief apparent differences from them are: the color is dark
lavender to plumbeous instead of brownish; ten folds on tail (from
front of vent) a somewhat larger series of splenial teeth; an in-
crease of 80 transverse folds above the highest in the other lot.
( See table for further data. )

That neither of these larvae is the young of Ichthyophis larutensis
sp. nov. herein described, may be deduced from the presence of
splenial teeth in the two types of larvae. These are absent in
larutensis which is a slenderer form, transforming at a distinctly
smaller size than the preceding larvae.

Ichthyophis paucidcntuhis sp. nov.

Type: U. S. N. M. No. 70671. Kapahiang, Sumatra; H. C. Kellers,
collector.

Diagnosis: A presumed medium-small species having a barely
discernible light streak on the side of body beginning at mouth
angle, discontinuous in numerous places; no trace of splenial teeth,
or of a ridge on inner side of jaw; eye moderately distinct through
a gray-white spot in the eye-area (in preser\ative); tentacular open-
ing lunate, relatively large, the tentacle conical, closer to eye than
to nostril; the accessory cusp is clearly visible on the front teeth of
maxillary and mandible; transverse folds, 393, 7 on tail; verte-
brae, 122.

50 The University Science Bulletin

Description of type: A medium-small, slender specimen, the
width in length, 34.5 times; head short, oval, the eyes dimly visible
through a gray-white circular mark; head width at first nuchal fold,
8 mm.; length of head, 11.5; distance between eyes (6 mm. on
curve) greater than snout length (4.9 mm.); eye to nostril, 3.2 mm.;
tentacle conical, the tentacular opening lunate, above rather than
behind tentacle, closer to eye (1.4 mm.) than to nostril (3 mm.);
distance from tip of snout to first nuchal groove, 10 mm.; to second,
13; to third, 17.5; first nuchal groove distinct on throat, not crossing
top of head; second visible below and on sides of neck; third indi-
cated laterally but barely traceable across neck.

Primary and secondary folds, 393, counted on dorsum (fewer,
380 counted ventrolaterally ) ; only a few anterior folds fail to
meet; other folds on venter meet at an angle, directed backwards;
each fold crosses the dorsum in a sinuous line and forms a convexity
directed forward on median line, its anterior position on dorsum
often from four to seven millimeters in advance of the ventral
angle; posteriorly a few folds cross the body and venter almost di-
rectly. The grooves do not cross venter except posteriorly, in front
of vent; four folds interrupted by the vent; no glandules * discernible
on sides of vent; tip of tail somewhat flattened on top, not or scarcely
compressed laterally.

Very small scales can be found in folds near the anterior part of
body, absent or scarce on the ventral region anteriorly; more pos-
teriorily the scales are larger and encircle body, about three im-
bricating rows in each fold.

Teeth: maxillary-premaxillary teeth, 25-27; vomeropalatine, 29-
30; mandibular, 23-24; no trace of splenial teeth nor of the inner
ridge or elevation that supports the splenials; most of the teeth
have an anterior accessory cusp well developed, the teeth hooked;
mandibular teeth larger than the maxillary-premaxillary series, and
these in turn distinctly larger than the vomeropalatines; tongue
large and rather thick, occupying all the space between the side
of jaws; choanae elongate, somewhat angulate on inner border,
separated by a distance slightly greater than twice the transverse
diameter of one.

Color in preservative: Plumbeous on dorsum and sides; \entrally
the shade is somewhat lighter; an indistinct, narrow, broken lateral
line of dull cream from angle of mouth to near level of front of vent;
a cream spot surrounds vent, widened at front end; tip of tail with

* These small glandules may be present only in males.

On the Caecilian and Related Species 51

a small cream spot; a gray-white spot surrounding and covering eye;
light whitish cream spots about tentacle and nostril.

Measurements in mm.: Total length, 294; tail, 6; head width, 8;
head length, 11.5; estimated body width, 8.5; width in length, 34.5
times; tail length in total length, 49 times.

Remarks: This species on the basis of the increased number of
vertebrae, and the lateral stripe, might seem to be closely allied to
Ichthyophis glutinostis. This is highly improbable since the entire
series of the splenial teeth is absent, and the jaw modified so that
the usual inner ridge that bears the splenials is likewise absent. It
probably is most closely related to /. nigroflavtis sp. nov.

The secondary cusp on the teeth is more or less indicated in many
species. In this form the size of the teeth makes the cusps more
evident. The number of the maxillary-premaxillary and vomero-
palatine teeth is, I believe, exceeded only in the largest known spe-
cies, Ichthyophis nigroflaviis sp. nov. and 7. malaharicus sp. nov. On
the other hand it would appear that the mandibular teeth are some-
what reduced in number.

The name paticidentuliis is derived from Latin paucus, few;
dentahis, toothed, in reference to the fact that there are only three
instead of four sets of teeth.

Ichthyophis monochrous (Bleeker)

Epicrium monochroum Bleeker, Nat. Tijdsclir. Ned. Ind., vol. 16, 1858, p. 188
(type locality Sinkawang west Borneo); Giinther, The Reptiles of British
India, 1864, p. 443 {part.) (refers a specimen from Singapore to this spe-
cies, specimen having 226 circular folds); Proc. Zool. Soc. London, 1872,
p. 591 (listing only).

Ichthyophis monochrous Peters, Monats. Akad. Berlin, 1879, p. 932 {part.);
Boulenger, Catalogue of the Batrachia Gradientia s. Caudata and Batrachia
Apoda in the collection of the British Museum, p. 91, pi. 4, fig. 1 {part.);
Boettger, Ber. Senckenberg. Nat. Ges., 1887, p. 50; Boulenger, The Fauna of
British India, including Ceylon and Burma. Reptilia and Batrachia, 1890,
517 {part.) (! Java, Borneo, Singapore, Sikhim and Western Ghats of India,
Malabar, Waghei Surat, India); Proc. Zool. Soc. London, 1895, pp. 403-404
{part.); ? de Elera, Catalogo Sistematico de toda la Fauna de Filipinas.
1895, p. 453; van Kampcn, Amphibien dcs Indischen Archipels.
Zoologische Ergcbni.sse eincr Reise in Niederliindisch Ost-Indien, herausge-
geben von Dr. Max Weber, 1907, p. 415 {part.); Nieden, Das Tierreich.
Lief. 37 1913 Gymnophiona, Berlin, p. 7 {part.); Bourret Les Batraciens de
rindochine (? date), pp. 138-139, fig. 15 {part.).

Type: British Museum Natural History No. 63.12.4.5 Sinkawang,
west Borneo.

Diagnosis: Vertebrae, 108; costal folds, aproximately 247; length
in width about 22-23 times; splenial teeth reduced, 4-4; four to five
scalerows in each fold, at least in posterior half of body; four an-
terior folds fail to meet (ventrally); transverse folds cross the body

The University Science Bulletin

Fig. 3. Ichthyophis monochrous (Bleeker). Type. B. M. N. H. No. 63.
12.4.5. Sinkawang, west Borneo. Left figure, dorsal view; right figure, ventral
view. Actual length of both figures, 232 mm.

On the Caecilian and Related Species

forming a strongly marked angle on venter; a narrow cream-colored
ring about eye, and a cream spot on vent; tail short, 3.8 mm., with
six folds.

Redescription of type: A moderately slender species, the length
232 mm.; width about 10 mm.; the width in length, approximately

Fig. 4. IcJithyophis monochrous (Bleeker) Type. Radiograph
showing 108 vertebrae

23 times; head oval in outline, the curved width between eyes, 6
mm.; width of head, 7 mm.; length of head, 9 mm.; snout length in
front of eyes, 4.2 mm.; eye visible, its diameter .4 mm.; tentacle
conical, situated near lip, closer to eye (1 mm.), than to nostril
(2.5 mm.), the tentacular opening lunate; nostril to lip, 1.4 mm.;
eye to lip, 1.15 mm.; length of head from first groove, 8.6 mm.; from

54 The Universit\' Science Bulletin

second groove to snout tip, 11.6 mm.; from third groove to snout
tip, 14.8 mm. ( measured on side ) ; the first nuchal groove is strongly-
distinct on the throat, dim above; second groove strong on throat
but not visible dorsally; third groove visible laterally, crossing above
neck as a typical costal groove, not completely crossing throat be-
low. This is preceded on dorsal surface by a short somewhat angu-
lar fold that terminates on the dorsal surface; the first four trans-
verse folds (following the third nuchal groove) do not meet on
venter; subsequent folds cross dorsum and sides in a sinuous line,
then bend backwards and meet at an angle on venter, at a point
four or more millimeters farther posterior than on dorsum. The
primaries and secondaries (perhaps tertiaries)* cannot be ordinarily
distinguished from each other.

There is a small but distinct tail bearing six transverse folds pos-
terior to anterior edge of vent, its length about 3.8 mm.; an area
about vent flattened; tail rounded above and somewhat compressed
laterally, terminally pointed; no trace of elevated lateral longitudinal
folds on body.

Skin strongly glandular throughout surface of body, and because
of the folding most of the larger glands lie recumbent in the skin,
very many having a length of more than one millimeter; each trans-
verse fold overlaps somewhat the deeper following fold; concealed
scales are present throughout surface of body, each fold covering
from one to five rows of transversely overlapping scales which en-
circle the body; the preceding several rows of scales, though sepa-
rated by tissue, actually overlap the anterior scale series of the
following fold; scalerows of each fold imbricated, the anterior over-
lapping the posterior.

The largest scales measure 1.8 mm. in greatest diameter. The
number of rows and the size of scales is reduced in the anterior part
of body. The vent is longitudinal with lateral grooves forming a
denticulate edge ( some 8 denticles on each side ) ; no gland is visible
on sides of vent. Teeth: maxillarv-premaxillarv, 25-25; vomeropala-
tine, 21-21; mandibular, 19-19; splenial 4-4. The tongue is broad,
rounded, partially covering the splenial teeth.

Color in preservative for one hundred years: Dark brown aboxe
and on head, the edge of each fold distinctly lighter, nearly fawn-
colored; eyes dark with whitish pupil surrounded by a narrow
cream-colored ring; a cream-colored area about vent occupying

* Some previous authors have suggested that there are two transverse folds to each
body seginent in Caecilians. This is certainly not true in all cases, since the vertebrae of
all specimens from S. E. Asia number less than half and often only a third of the number
of transverse folds. Thus it must appear that numerous vertebrae have more than two
transverse folds.

On the Caecilian and Related Species 55

space between the interrupted folds; area about nostril dull cream;
grooves on throat lighter than surrounding color. In life the color
is said to have been violet-brown.

Measurements in mm.: Total length 226 (Bleeker's measurement
232) width of body, 10; width of head, 7; head length 9; largest
scales, 2 mm. in diameter; width in body length approximately
22-23 times.*

Remarks: The specimen has been somewhat dehydrated and the
back part of the body has had the folds loosened.

Ichthyopliis singaporensis ** sp. nov.

Ichtlujophis glutinosiis, var. Cantor, Journ. Asiat. Soc. Bengal, vol. 16, p. 1058,
and reprint in "Miscellaneous papers relating to Indo-China for Straits
Branch of Royal Asiat. Soc," vol. 2, Trvibner and Co., London, 1886, p. ?;
Boulenger, Catalogue of the Batrachia Gradientia s. Caudata and Batrachia
Apoda in the Collection of the British Museum, 2nd ed., 1882, p. 91.

Type: British Museum No. RR. 1959.1, 2.43. Singapore.***

Diagnosis: A small species having 267 primary and secondary
folds about the body, 7 confined to short tail; maxillary-premaxillary
teeth, 24-24; vomeropalatine, 20-20; mandibular, 20-20; splenial,
3-3. Orbit completely surrounded by bone; the tentacular opening
in maxillary bone close to lip, nearer to eye than to nostril; sepa-
rated from orbit by bone for a distance greater than the diameter
of the opening. Total length of type, 243 mm. Tail length in total
length 54 times; body width in total length 22 times. Vertebrae 111.

Description of type: (External characters of the head missing;
its general characteristics and size may be derived from the skull
measurements given). Tentacular opening near lip, closer to eye
than to nostril; estimated head length, 10 mm.; estimated width of
head, 7.5; first nuchal groove can be traced around head, most dis-
tinct below; second nuchal groove distinct below, not reaching
dorsal surface of neck; third groove seemingly distinct dorsally and
more or less distinct ventrally.

Scales present in first transverse folds, large, and in one or two
continuous rows about body (except median point on venter).
These continue throughout body, the rows growing more numerous
in each fold until posteriorly there may be 7 or 8 rows in a single
fold, the fold 3 to 3.2 mm. wide, the scales imbricating and over-
lapping laterally, as well as passing entirely around body. Many

* The partial dehydration of the specimen has changed the measurements.

** Singapore + ensis = place, locality, country.

*** The skin of the head had heen loosened from about the skull to expose the skull
while it was still attached to the vertebral column. The skull has now been placed in a
small glass container attached to the type specimen.

The University Science Bulletin

Fig. 5. Ichthyophis singaporensis sp. nov. Type. Radiograph
showing 111 vertebrae.

scales in posterior folds visible through the skin in this .specimen
(probably due to state of preservation); elongate recumbent glands
opening between the folds. Transverse folds, dorsolateral count,
267, with 7 confined to tail; first three chevron-shaped folds on
dorsum above pharynx occurring between the two posterior nuchal
grooves, do not reach ventral part, but have well-developed scales;
the fourth and succeeding folds completely surround the body,
ventrally forming an angle, the transverse grooves seemingly not
traceable on ventral parts until a short distance in front of vent
where the folds and grooves cross venter without forming an angle;

On the Caecilian and Related Species

vent longitudinal, interrupting four folds. Vertebrae 111. (Tooth
formula in diagnosis. )

Color in preservative: Dull dark brown; if a cream spot occurs
at vent it has now been discolored; a rather broad cream line bor-
ders the lower edge of each lower jaw, a darker Hne separating this
from the lip; no cream spots discernible at eye, tentacle, or nostril.

Measurements in mm.: Total length (estimated for missing head
based on skull length), 243; tail, 4.5; estimated width of head, 7.5;
length of head, 10; body width, 11; width in length, 22 times; tail
in total length, 54 times.

'*'**^^^*5:KRRr5j5c^rcSisi'

Fig. 6. Ichthyophis singaporensis sp. nov. Type. Fig. 1, dorsal view of
skull; fig. 2, lateral view of skull; fig. 3, lower jaw, much enlarged.

Remarks: Figures of the skull show most of the important rela-
tions of the bones of the skull. One of the most significant features
is that the orbit is closed completely and widely separated from the
tentacular opening in the maxillary.

Figure 28,* Epicrium ^lutinosum given by Wiedersheim offering
a lateral view of the skull shows 'the tentacular canal continuous
with and forming a forward and downward extension of the orbital
cavity. I am uncertain of the source of the specimen figured by
Wiedersheim and cannot be certain that it even represents ^lutino-
siis as here considered. However it suggests the possibility that two
genera are involved.

It is with some reluctance that I have used this ancient specimen,
now badly mutilated, as a type specimen. The characters however,
point to a distinct species. It came from a garden on Singapore

* Die Anatomip der Gymnonhionen. Tena 1879, Plates: 2, fig. 23; 3, figs. 25, 26, 28. 29,
31; 4. figs. 35, 38 to 42, 43; 6, figs. 62, 69, 70; 7, figs. 75, 80; 9, fig. 88.

58 The University Science Bulletin

Island and to the best of my knowledge is the only specimen that has
been reported as ha\inc; been captnred there despite the interven-
tion of at least one hundred ele\'en years. I failed to find it during
a stay of some weeks on the Island. If the specimen actually came
from some other locality it is no less remarkable since it has not
been reported elsewhere.

That the species may be related to IcJitliyopJiis monochroiis is
shown in the body proportions, and the reduced splenial teeth.
They differ however in the development of the scales, and the num-
ber of transverse folds is somewhat larger in singaporensis. The
matter of relationships cannot be decided until the skull characters
of 7. monochroiis are known.

Ichthijophis duJitensis * sp. nov.

Jchthtfophis monoclirous Boulenger, Proc. Zool. Soc, London, 1892, p. 508
(Mt. Dulit, Borneo).

Type: British Museum of Natural History, No. 92.6.3.23, Mount
Dulit, Sarawak, Borneo; elevation above 2000 feet; collected by Mr.
Charles Hose, autumn 1891.

Diag,nosis: A mountain species, 235 mm. in length, having 114
vertebrae and 313 transverse folds; splenial teeth reduced (4-4);
the body width in length about 29 times; head of lighter color than
body; a large cream spot on chin and throat; pharyngeal region
darker than body or head.

Description of the type: A moderately slender species, the width
in length approximately 29 times; head width at first nuchal groove,
8 mm.; the length of head, 11 mm.; width between eyes, 5.15 mm.;
length of snout in front of eyes, 4.2 mm.; eye small with a white
pupil surrounded by a black iris, slightly elevated on surface of
head, its diameter, .9 mm.; head length from snout-tip to first nuchal
groove, 10.5 mm.; to second groove, 13.2 mm.; to third, 17 mm.;
tentacle conical, the opening lunate, the area about opening swollen,
closer to eye (1.5 mm.) than to nostril (2.5 mm.); nostril and eye
about equidistant from lip, the distance between them 3.4 mm.; the
first nuchal groove is distinct on throat and sides of head but very
dim or absent on dorsal part; second groove distinct on throat and
discernible on side of neck, but not on the dorsal region; the third
groove can be discerned on the sides of neck only.

The primary and secondary transverse folds, 313 (dorsal count;
304 ventral count ) ; all grooves and folds appear to pass completely
around the body throughout the length, tending to turn backward

* (Mt.) Dulit -f- cnsis (Latin) = place, locality, country.

On the Caecilian and Related Species

Fig. 7. Iclithyophis cluliteus-is sp. iion . Type. B. M. N. H. No. 92.6.3. 2o,
Mt. Dulit, Sarawak, Bornto, ele\ . ahoM- 2()()() ft. Left figure dorsal view,
right figure, ventral view. Aetual k-ngtii, 2.35 iinn. Note tliat the head ot the
left figure and the ventral side of tlu- posterior part of the bod\ are light struck.
The white spot on the throat is natural.

60 The University Science Bulletin

and meet at an angle on \entral snrface. The angles widen pos-
teriorly and folds in front of vent pass almost directly across; seven
folds on the tail (counted from front of vent); vent somewhat cir-
cular with a depression and groove following; a pair of tiny eleva-
tions on each side of vent suggesting glands.

Fig. 8. Ichthyophis duUtensis sp. nov. Type. Radiograph
showing 114 vertebrae.

Scales present throughout body, those on anterior part much
reduced in number and size. In posterior part there may be as
many as four or five rows of scales in each fold, which partially
overlap the scales of the following fold.

Teeth: maxillary-premaxillary, 18-18; vomerpalatine, 23-23;
mandibular, 20-20; splenial, 4-4.

On the Caecilian and Related Species 61

The tongue is rounded, perhaps not covering the splenials com-
pletely; choanae rather large, the diameter of one ( .76 mm. ) in the
space between them (2 mm.), 2.63 times; the tail length in total
length, about 42 times.

Color in preservative: Head olive brown; pharyngeal region very
dark black-brown contrasting with both body and head; brown gen-
erally over body, the front part of the transverse folds darker than
the remainder so that fine lines of dark brown appear to be alter-
nating with fine lines of tan; a cream spot surrounding the vent and
the depression following; a cream spot about tentacle and nostril;
a small cream ring about eye. The large cream spot on throat
somewhat broken up by brownish lines in the nuchal grooves.

Measurements in mm.: Snout to vent, 235; tail, 5.6; head length,
11; head width, 8; average body width, 8.

Remarks: This species, which geographically is close to the iyge
locality of Ichthyophis monochrous is clearly differentiated from the
latter species by the much larger number of primary and secondary
folds (313 instead of 247), the presence of a cream spot on the
throat, seemingly a different type of vent, a larger head, and six
more vertebrae.

They agree in having a reduced series of splenial teeth, and the
body folds have the same type of ventral angle.

From I. weberi it differs in having a series of splenial teeth; from
7. javanicus in being less slender, in having a reduced series of
splenial teeth (4-4 instead of 12-12) and fewer transverse folds
(313 instead of 351); from I. gJandulosiis in lacking the lateral
ridges, in having folds and grooves encircle the body, more trans-
verse folds (313 instead of 273), and a reduced series of splenial
teeth.

Jchthijoph's peninsuJaris * sp. no\'.

lclithy()))liis monochrous Boulenger, Ciituloguc of the Batracliia Gradicntia s.
Caudata and Batrachia Apoda in the Collection of the British Museum, .'Znd
Ed., 1882, p. 91, pi. 4, fig. 1 (paii. Malabar): Fauna of British India, in-
cluding Ceylon and Burma. Rcplilia and Batrachia, 1890, p. ,517 {part.);
Ramaswami, Current Science, Bangalore, vol. 16, 1, 1947, pp. 8 to 10
(? part.).

Type: Brit. Mus. Nat. Hist., No. 82.12.12.6; Malabar, India, R. H.
Beddome collector.

Diagnosis: A large species, with a broad, relatively short head;
the eye invisible covered with a raised circular white spot; the ten-
tacle nearer the eye-spot tlian to tlic rostril; tail long, its length

* Peninsula (Latin) referring to peninsular India.

The University Science Bulletin

Fig. 9. Ichthyophis pemnsidarls sp. nov. Type. B. M. N. H. No. 82.12.12.6,
Malabar, India. Actual length, 330 mm.

On the Caecilian and Related Species

in total length nearly 22 times; the body width in length about 22
tacle nearer the eye-spot than to the nostril; tail rather long, its
length in total length nearly 22 times; the body width in length about
22 times; transverse folds on dorsum, 363; folds on tail, 18; vertebrae,
116; splenial teeth, 4-4. Ventral surface very light, probably cream
colored in life; a well-defined cream spot at vent.

Fig. 10. Ichtliyophis peninsularis sp. nov. Type. B. M. N. H.
No. 82.12.12.6. Radiograph showing 116 vertebrae.

Description of type: Width of head ( 14.4 mm. ) only a little less
than length of head (16 mm.); distance between eye-spot, 7.1 mm.
measured on the curve, greater than length of snout in front of eyes
(6.1 mm.); tentacle withdrawn, the tentacular opening circular;

64 The University Science Bulletin

tentacle opens near lip between eye and nostril, closer to eye-spot
(2.5 mm.) than to nostril (4 mm.); eye-spot and nostril equidistant
from lip, the distance between them 5 mm. The anterior nuchal
groove encircles the head; second fold well defined on neck, and
somewhat on sides of neck; third fold more or less distinct except
in the median ventral area; snout tip to first groove, 14 mm.; to
second, 19 mm.; to third, 24 mm.; six anterior folds fail to meet be-
low on the median line; the folds can be seen because of the distri-
bution of the glands but the grooves themselves seemingly do not
cross the venter except in the posterior part of the body; folds
counted on dorsal surface, 363; on ventral surface, 363, forming an
obtuse angle directed backwards except in the posterior part of
body where they pass directly across; tail somewhat compressed
laterally and pointed terminally; the vent interrupts eight folds; a
total of 18 folds on tail from front of vent.

Scales are present throughout body, anteriorly somewhat smaller
and sparse or absent ventrally; posteriorly there are from three to
five transverse rows in each fold extending around the body; ver-
tebrae, 116.

Teeth: maxillary-premaxillary, 24-25; vomeropalatine, 25-26;
mandibular, 25-24; splenial, 4-4. The maxillary teedi are larger
than the vomeropalatine series, but are somewhat smaller than the
mandibular or at least the anterior ones.

Coloration in preservative: Above, grayish lavender growing
lighter laterally and ventrally; imderside with some pigment but
light, perhaps cream-colored in life. Head somewhat grayish
brown; the eye-spots whitish; small cream areas about tentacle and
nostril.

Measurements in mm.: Total length, 330; tail, 15.2; width of body,
14.8; head width, 14.4; head length, 16.

Variation: A second specimen available agrees with the preceding
in general; the following data are given for comparison: length, 258.
tail, 11.8, width of body, 13, width in length, 20, head width, 11.8,
head length, 14; transverse folds (dorsal) 366, tail folds. 18. Teeth;
maxillary-premaxillary teeth, 20-20; vomeropalatine, 19-19; mandib-
ular, 15-15; splenial, 3-4. The specimen is one recently collected
and is darker than the preceding.

A Berlin specimen from Malabar is young, measuring 222 mm.
in length. The transverse folds number 357, the vertebrae, 116. and
the splenial teeth, 4-4. The specimen is light lavender to violet
above; below, the chin and neck are cream-white, and the remainder

On the Caecilian and Related Species

of the venter is dull cream. The head is lighter in color than the
remainder of the dorsum.

Remarks: It is not impossible that the type is somewhat faded
since it has been in preservatives considerably more than half a
century. It is, however, in relatively good condition except that
exploration of the mouth has injured the tongue and the skull is
broken in the palatal area.

Fig. 11. Ichthyophis peninsularis sp. nov. Type,
ot the subcaudal region. Enlarged.

Characteristics

Ichtliyophis subterrcstris * sp. nov.

Type: Chicago Natural History Museum No. 73927, Travancore-
Cochin, plains, India.

Paratype: Stanford Mus. No. 2118, from Kottayam, Travancore,
India.

Diagnosis: Dark violet-lavender above and dark lavender-brown
venter; a light area about nostril, but not about tentacle or vent;
tentacular opening circular; mandibular teeth very large, approxi-
mately three times the size of teeth in maxillary series; splenial

* Sub (Latin) under + terrestris (Latin) = of the earth.

3—8920

66 The University Science Bulletin

teeth, 5-5; 364 folds, 18 on tail. Total length, 260; tail, 11.7; tail
length in total length, 22.2 times; body width in body length ap-
proximately 21 times.

Description of type: Head somewhat elevated posteriorly, the
width at first nuchal groove 11 mm.; the length of head, 14 mm.;
distance between eyes on curve (6-8 mm.) greater than length of
snout ( 5.35 mm. ) ; tentacle withdrawn, not attached to the edge
of opening; the tentacular opening close to lip, circular; tentacle
close to lip, much closer to eye (1.8 mm.) than to nostril (3 mm.);
distance between eye and nostril, 4.3 mm. Distance from tip of
snout to first nuchal groove, 13 mm.; to second, 16.8 mm.; to third,
20.2 mm. (measurements made on side of head).

Primary and secondary folds together 364, of which 18 are on
tail; these cross the body, and form angles ventrally, except pos-
teriorly; first four or five failing to meet on throat, eight interrupted
by vent; anterior folds curve forward across neck; the grooves are
not complete on venter except in the posterior part of the body;
small scales are present from the first fold, both dorsally and
ventrally, only one or two scalerows usually present; farther back
scales increase in size and number the posterior folds having four
or five rows, the first row of each series smallest, the one preceding
the last largest, all imbricating.

Teeth: maxillary-premaxillary, 21-21; vomeropalatine, 21-21;
mandibular 17-17; splenial, 5-5. The size of the teeth is such that
the tiny notch near the outer anterior face of the tooth can be seen.
The tongue is large, completely covering the splenial teeth, the an-
terior part broadly arrow-shaped and somewhat elevated. It seems
probable that some of the posterior splenial teeth are missing since
the paratype has a somewhat larger number.

Tail laterally compressed, the terminal part without folds (true
in practically all species); a pair of darker areas along side of vent
perhaps representing the position of special glands; choanae some-
what angular on their inner face, separated from each other by
a distance equal to three times the transverse diameter of one.

Color in preservative: Dark violet-lavender above, becoming
brownish-lavender on venter; slightly lighter about eye; however,
a few small rounded flecks, presumable glands are present close to
eye; a dull cream area about nostril; area about vent lighter than
surroundings (perhaps originally cream).

Measurements in mm.: Total length, 260; tail, 11.7; tail in total
length, 22.2 times; body width, 12.4; head width, 11; head length, 14.

On the Caecilian and Related Species 67

Remarks: The absence of calcium in the body was discovered on
examining an X-ray picture of the specimen. There is no trace of
the skeleton to be discerned. This absence is probably caused by
the method of preservation. It is a relatively recent specimen and
whatever the chemical, so effective in dissolving calcium, it seem-
ingly has not affected the color and texture of the other tissues.

The paratype agrees with the type in very considerable detail
except that the skeleton is intact and there are three more splenial
teeth on each side. The characteristics of the scales and their dis-
tribution are those of the type. There are 356 folds, 16 on tail.

The maxillary-premaxillary teeth are about 27-27; the vomeropala-
tines, 25-28; the mandibular, 20-20; and splenials, 8-8. The length
is 295 mm. The skin on one side of the head has been removed dis-
closing the orbit and the tentacular groove. The latter is completely
separated from the orbit occurs in singaporensis, as is indicated in
fig. 6. The shape of the tentacular opening is somewhat more
rectangular. The increase in number of teeth may be due to age.

This species is seemingly distinguished from other species of the
Indian Peninsula by the following combination of characters: the
shape of the tentacular opening; the elongate tail with the high num-
ber of folds; the tail length being contained in total length only 22,2
times; the enlarged mandibular teeth. The color of the venter
separates it from peninsularis and the body proportions seem to be
somewhat different.

Idithijophis homhaijcnsis * sp. no\ .

Type: British Mus. Nat. Hist. No. 86.6.11.1, Waghii Surrat, Bom-
bay, India; Gleadon, collector.

Diagnosis: A large species (390 mm.) having 386 transverse
folds, 14 confined to tail; splenial teeth 9-9; body width in length, 26
times; vertebrae, 121; color in preservative, dark brown above, some-
what lighter brown below; tail relatively long contained in total
length about 25 times; tongue tending to cover the splenial teeth;
mandibular teeth much larger than maxillary-premaxillary series;
vomeropalatine teeth relatively small, scarcely extending through
the thick gums.

Description of type: Head proportionally small, its width at first
nuchal fold 10 mm., the length 15 mm.; eye distinct, the lens a
pearly sphere surrounded by a narrow black rim of iris and these in
turn surrounded by a light ring on skin wider above eye, very nar-

* Bombay + ensis = place, locality, country.

The Uninersity Science Bulletin

row below; tentacular opening somewhat rounded, the tentacle a
fine slender cone seemingly not attached laterally to the opening;
tentacular opening about half a millimeter from lip, closer to eye
(2.3 mm.) than to nostril (4 mm.); eye from nostril, 5.5 mm.; eye
from lip, 1.9 mm., nostril from lip, 1.8 mm.; distance between eyes
(on curve), 8 mm.; length of snout, 7 mm.; distance between nos-
trils, 4 mm. First nuchal fold well visible above on sides and on un-
derside of neck, preceded by a distinct strongly curving fold on
back of head beginning above mouth angle, not visible on sides or

Fig. 12. Ichthyophis bombayensis sp. now Type. B. M. N. H. No. 86.6.11.1.
Waghii Surrat, Bombay, India. Actual length, 390 mm.

below; second nuchal fold not discernible on sides or above, dis-
tinct below; third nuchal groove strong on sides, less so on dorsum,
partly crossing venter. Snout to first nuchal groove (lateral meas-
urement), 15 mm.; to second, 20 mm.; to third, 27 mm.

Transverse folds on body. 386; 13-14, folds confined to tail; tail
rather strongly compressed laterally; transverse folds make a median
forward curve dorsally, then on sides tend to turn back and form a
median ventral angle at a point five to seven millimeters behind its
most anterior point above. Scales are present throughout, anteriorly

On the Caecilian and Related Species 69

usually only a single row in each fold, more posteriorly there may
be three imbricating rows, the middle one much the largest in
each fold.

Teeth: maxillary -premaxillary, 23-22; vomeropalatine, 24-24;
mandibular, 20-20; splenial, 9-9.

Color in preservative: Above dark brown the posterior edge of
the folds a little lighter; top of head somewhat mottled with darker
brown; venter lighter brown; a cream spot surrounding vent. A
small light area around eye; a cream spot below tentacle and one
about nostril; lips and an area at mouth-angle cream.

Measurements in mm.: Total length, 390; tail, 15.2; head width,
10; head length, 15; width, 15; width in length, 26 times; tail
length in total length, 25.6.

Remarks: The increased number of vertebrae, the coloration, the
enlarged mandibular teeth and the reduction of the size of the
vomeropalatine teeth seem to separate this from more southern In-
dian forms. The specimen is a male.

Ichthyophis mindanaoensis * sp. uo\.

Ichthyophis monochrotis Inger, Fieldiana Zool., vol. 33, no. 4, 1954, pp. 207-209
( part. ) ; Todaya on Mount Apo, Davao Province, Mindanao, P. I.

Type: Chicago Natural History Museum No. 50958, Todaya, Mt.
Apo, Davao, Mindanao, P. I., 2800 feet elevation, H. Hoogstral, col-
lector.

Poratype: C. N. H. M. No. 50957, Mt. McKinley, Davao, Min-
danao, P. I.

Dkip,nosis: A medium-sized species, largest known specimen, 276
mm. (283 fide Inger). Vertebrae, 110-116; transverse folds, 308-
317; 322-341; body width in length approximately 28; tail length
in total length approximately 43 times; splenial teeth, 8-8 to 11-11.
Single gill opening on each side in larvae.

Transformation occurs in larvae after a length of 238 mm. has been
reached, the large larvae retaining the lateral line system clearly
defined on head. Differs from glandulosiis (from Basilan, P. I.) in
having 8 to 14 more \ertcbrae, more (30 or more than highest
count) transverse folds. The choanal openings are at least one half
larger.

Description of type: Head rounded, oval; snout projecting .some-
what; eyes visible, covered with .skin that forms a slightly raised
circular area, dark in the middle and ringed with gray; width of

Mindanao + ensis (Latin) = place, country of.

The University Science Bulletin

Fig. 13. Ichthyophis rnindanaoeiisis sp. nov. Type. Chicago N. H. M.
No. 50958. Todaya, Mt. Apo, Davao, Mindanao, P. I. 2800 ft. elev. Upper
figure, dorsal view; lower figure, ventral view. Actual length, 276 mm.

On the Caecilian and Related Species

head at first nuchal groove, 9.3 mm.; length of head, 14 mm.; tip of
snout to first nuchal groove, 12 mm.; to second, 15 mm.; to third, 20
mm.; distance between eyes measured on curve (6 mm.) longer
than snout (5.1 mm.); the first nuchal groove distinct on throat can
he traced across top of head; second, distinct on throat and on side
of neck for a short distance; third, visible across dorsum and on
sides, scarcely or not discernible on throat. Tentacular opening
lunate, near edge of lip, much closer to eye (1.5 mm.) than to nos-

FiG. 14. IdUhyophis mindanaoensis sp. nov. Upper fig., type; lower fig.,
paratype. C. M. \. II. No. .50951, Mt. McKinley, Davao, Mindanao, P. I.

72 The University Science Bulletin

tril (3.6 mm.); distance between eye and nostril, 4.5 mm.; nostril
closer to lip than eye. Transverse folds, 305, six confined to tail;
folds and grooves complete across dorsmn, the grooves failing to
cross the middle part of venter except on the latter fourth or fifth of
body. The vent interrupts three folds.

Choanal openings large, the distance between the edges of the
choanae, 2 mm.

Scales present; anteriorly sparse, small, usually in a single row
on dorsum in each fold; more posteriorly the scales increase in size
and number until there are three or four rows surrounding the body
in each fold. Vertebrae, 111.

Teeth: maxillary-premaxillary, 25-26; vomeropalatine, 24-24; man-
dibular, 17-18; splenial, 8-8.

Color in preservative: Above plumbeous-violet to brownish, the
venter a slightly lighter shade; a cream spot covering vent, one
about tentacle and one around nostril; a gray ring surrounds eye
connecting with a gray area lying between eye and tentacle; some
fine light areas under throat on midline. Lips lighter than sides of
head; under clear liquid the body segments can be discerned along
the median ventral line, as indicated by slightly darker transverse
marks.

Measurements in mm.: Total length, 276; tail, 6.4; width of body
approximately, 9.8; head width, 9.3; head length, 14.

Remarks: There is a second adult specimen in the collection from
Mt. McKinley, Mindanao, P. I., taken at an elevation of 3000 feet,
that may belong in this species. The specimen is 7 mm. longer
than the type of mindanaoensis and has 113 vertebrae; there are 308
folds, 9 of which are confined to the tail. The scales are scattered,
or absent and small when present on the anterior half of the body;
at first only a single row present but in the posterior folds there are
three rows, transversely overlapping, and imbricating.

Larvae: There is a series of larvae taken at Todaya, Mt. Apo, 2800
feet elevation, "to be found in rivers and on the ground during rain,"
that presumably represent this species. These specimens are re-
corded in the accompanying table.

The neuromast (lateral line) system is retained and is especially
distinct in all but the largest larvae. It would appear that this is an
adaptation for life in deeper waters such as rivers as opposed to
brooks. In most of the species studied only the small and presumed
very young larvae showed the organs clearly. The larvae are large
at the time of transformation (238 mm.).

On the Caeciliax and Related Species

Fig. 15. IcJithyophis miudaiuiocnsis sp. nov. Paratype larva. C. M. N. H.
No. 50970, topotype. Actual length, 220 mm. Showing the details of the
neuromast system in large lar\'ae. Actual length, 220 mm.

Table of data on larval specimens presumed to be young of Ichthyophis

mindanaoensis sp. nov.

Number

Length

Vertebrae

Body folds

50966

98
120
150
- 156
191
220
220
224
231
236
236
237
238

50967

50973

50969

50968

50961

50960

116
116
114
114
116
116
114
114

320

50970

327

50963

344

50964

330

50975

341

50971

324

50972

302

50976

334

Dr. Inger, with his characteristic graciousness, has kindly per-
mitted me to study these adult specimens from Mindanao and the
larval series which may be of the same species. Our findings are,

74 The University SciexNCe Bulletin

of course, quite different. I cannot agree that these specimens are
conspecific with IcJitliyopJiis monochrous Bleeker, Ichthijophis
ghndujos'us Taylor and Iclifliyopliis weberi or with any one of
them.

Dr. Inger's diagnosis of /. monocliroiis reads as follows: "Body
cylindrical or slightly depressed, elongate; numerous annuli around
the body; eye small, covered with skin; tail short, pointed; anal
opening a longitudinal slit." A half or more of the known caecilians
might easily fit this diagnosis and certainly the above mentioned
species would thus be included in IclitJiyophis monochrous.

His description is perhaps a composite made up from his speci-
mens and from the literature since his \ariation in annular rings is
somewhat greater than is shown in the Mindanao material.

Dr. Inger states that there is no tentacular aperture but he has
overlooked this in the oldest larvae where a pit or aperture ap-
pears in the eye area; he also states that there are no spiracles, but
these are certainly present in 14 of the larvae studied and are prob-
ably present in the entire 18 mentioned.

Ichthyophis ghindiilosus Taylor

IcJithyophis glandulosus Taylor, Philippine Journ. Sci., vol. 21, no. 3, Sept.

1922, pp. 516-517, pi. 3, figs. 4, 5 (tvpe locality, Abung Abung, Basilan I.,

P. I.).
Ichthyopliis monochrous Inger, Fieldiana Zool., vol. 33, no. 4, July 23, 1954

( parts ) .
Ichthyophis Taylor, Distribution of life in the Philippines, Bureau of Science,

Monograph, no. 21, Manila, 1928, p. 219 (Zamboanga).

Diagnosis: A medium sized species (largest known specimen 250
mm.); 273 transverse folds and grooves, meeting on dorsum in neck
region and in the latter two fifths of body, rarely meeting on venter
except in latter part of body; inner mandibular splenial teeth 11-11;
tongue somewhat pointed anteriorly rather than rounded; distinct
dorsolateral ridges. Vertebrae, 102.

Redescription of type: Cal. Acad. Sci. No. 60073 Abung Abung,
Cotobato, Mindanao, P. I. Moderately slender, the length, 250 mm.,
the width of body, 11 mm., the width in length about 23 times;
eyes very dim, but visible, the distance between them (measured
on the curve), 5.8 mm.; snout length in front of eyes, 4.5 mm.;
width of head at first groove, 9.3 mm., length of head, 12.5 mm.;
eye slightly elevated, its diameter, .95 mm.; tentacle not exserted, the
opening vertically lunate, situated very close to edge of lip, 1.3
mm. from eye, and 3.5 mm. from nostril; nostril and eye each about
1.2 mm. from lip; tip of snout to first grooxe, 11.5 mm., to second, 15

On the Caecilian and Related Species 75

mm., to third groove, 17.2 mm. (these three measurements are made
on the side of head and neck ) .

First nuchal groove deeply marked on throat and on sides, less
distinct dorsally and not, or but dimly crossing dorsal area; third
groove distinct laterally but only faintly indicated on ventral and
dorsal surfaces. Grooves following do not meet on the ventral part
of body except the 22 preceding vent; the first 22 meet dorsally, and
the terminal 70 grooves meet. The folds do not meet dorsally
throughout much of body where grooves fail to meet (or their
meeting not discernible); four folds interrupted by \ent.

Scales present posteriorly, three or four rows of scales in each
fold. Farther forward there may be only two rows of very small
scales; and anteriorly they seem to be entirely absent in first 40 or
50 folds, perhaps more. When present scales of one fold tend to
overlie partially those in the following fold, although separated by
tissue.

There are 273 folds altogether, six on tail; posteriorly, where they
can be seen to meet, they form a slight angle directed posteriorly; a
pair of sharply defined lateral folds or ridges extend throughout
body to within 2.5 centimeters of the tail tip.

Skin glands are present throughout the body and many can be
seen through the skin; the largest ones seem to have their openings
along the edges of the grooves and tend to lie recumbent in rows
below the scales. There are two swollen areas on each side of the
anterior part of the vent suggesting special glands. Vent longi-
tudinal, the edges denticulate; the terminal part of tail lacking
folds; sides of tail slightly compressed, the tip somewhat pointed.

Teeth: maxillary-premaxillary, 23-23; vomeropalatine, 24-24;
mandibular, 21-21; splenial, 11-11. The tongue pointed rather than
rounded, partially covering the splenial teeth; choanae small, the
transverse diameter about .35 mm., contained into distance between
them about 6 times.

Color in life: "Deep lavender to slate; more olive than lavender
on venter. In formalin the specimen is brownish-lavender with
scattered deep purple markings." After 38 years in alcohol the speci-
men brownish, with a slightly lighter brown \enter, the purple
marks indicated. A cream area around vent.

Measurements in mm.: Total length, 250; tail from front of vent,
6.2; head width, 9.3; head length, 12.5; width of body (average)
about 11; width in length approximately 22.7 times.

The University Science Bulletin

Fig. 16. Ichthyophis slandidosis Taylor. Type. Cal. A. S. No. 60073, Abung
Abung, Cotabato, Mindanao, P. I. Actual length, 250 mm.

On the Caecilian and Related Species 77

Variation: There are three paratypes; one measuring 165 mm. was
(in life) a deep slate-purple; the lateral folds are present in all.
The posterior part of the head is less widened than in the adult.

The specimens were obtained under fallen logs in moist situa-
tions along a small forest stream in southern Basilan. The species
can be separated from Ichthyophis weberi by the long series of
splenial teeth (11-11), none in weberi, and the low number of
folds about body (273 compared to 324 in weberi), and the fact
that the grooves do not cross the dorsum throughout much of the
body. From monochrous it may be separated, by the failure of the
grooves to meet on the mid-ventral line in much of the body, in
the longer series of splenial teeth and in the presumed greater num-
ber of body folds (273 compared to 247 in the type of monochrous),
and fewer scales in each fold.

Remarks: A specimen of Ichthyophis was taken near the city of
Zamboanga, in the Province of Zamboanga, Mindanao, Philippine
Islands in 1923. I was unable to make a study of the specimen but
it was tentatively identified as "Ichthyophis ( ? ) glandulosus Taylor."
On my return to Manila in 1957 I learned that this specimen as well
as the type of Ichthyophis weberi had been destroyed in the final
battle fought in Manila of World War II. A last stand was made
by the Japanese in that section of the building where I formerly
had my office and which housed the extensive herpetological and
ichthyological collections. All collections were destroyed. There
is no certainty that the species was glandulosus. The paratype
material is not now available to me for study.

Ichthyophis javanicus* sp. nov.

Type: British Museum of Natural History, No. 80.5.7.3. Col-
lected, "Java" the exact locality not known.

Diagnosis: A slender species, the greatest known length, 210
mm., the body width, 6 mm.; the tail length, 4.5 mm.; width in
length, 35 times; splenial teeth, 12-12; vertebrae, 115; primary and
secondary transverse folds, 351 dorsal, — 348 ventral count; ten
folds on tail, five interrupted by longitudinal vent; tentacle nearer
to eye than to nostril.

Description of type: A slender species the head slightly elevated,
the height of head about 2.4 mm.; width between eyes, 4.5 mm.;
length of snout anterior to eyes, 3.5 mm.; width of head at first
nuchal groove, 6 mm.; length of head, 9.4 mm.; eye very distinct,

* Java -f- icus = belonging to, pertaining to.

The Uni\tersity Science Bulletin

the pupil white, the iris black with a very small cream area in front
of eye; an incomplete ring of whitish flecks about eye; tentacle
close to edge of lip, the tentacle conical seemingly somewhat
rounded at tip, the tentacular opening lunate; tentacle nearer to
eye (1 mm.) than to the nostril (2.3 mm.). First nuchal groove
clearly indicated only on sides of head (or neck) but on throat
vaguely indicated; second groove vaguely indicated on sides and

Fig. 17. Ichthyophis javanicus sp. nov. Type. B. M. N. H. No. 80.5.7.3.
"Java." Actual length, 210 mm.

On the Caecilian and Related Species

underside of neck; third groove no different in appearance from a
primary costal groove; pharyngeal region not wider than head; tip
of snout to first groove, 9 mm.; to second groove, 11.7 mm.; to third,
13.7 mm.

Approximately 351 primary and secondary folds surrounding
body, meeting on the median ventral line at an angle which be-
comes broader posteriorly; ten folds on tail of which five are inter-
rupted by vent; the transverse grooves likewise appear to sur-
round the body; scales on anterior part of body absent or
very small and scattered; posteriorly the scales form transverse
rows, the scales overlapping, usually only one or two rows present

Fig. 18. Ichthyophis javanicus sp. nov. Type. Radiograph
showing 115 vertebrae.

80 The University Science Bulletin

in each fold, the scales varying much in size. There are no dorso-
lateral ridges evident.

Teeth: maxillary -premaxillary, 19-19; vomeropalatine, 18-19; man-
dibular, 19-19; splenial, 12-12; the mandibular teeth largest, the
splenials smallest; the tongue is somewhat narrowed anteriorly not
covering the vomerine teeth.

Color in preservative: Entire body brown, the ventral surface
only slightly lighter in shade; a cream-colored area about tentacle
and nostril; a cream spot on vent widened anteriorly; the anterior
edge of each fold slightly darker than remaining part.

Measurements in mm.: Total length, 210; tail length, 4.5; head
width, 6; head length, 9.4; eye to nostril, 3.

Remarks: This species is easily differentiated from Ichthyophis
glandulosiis in having 78 more primary-secondary folds; from
/. monochrous in having 104 more folds, and 115 rather than 108
vertebrae; from 7. weberi in having a well-developed series of
splenial teeth in the lower jaw ( none in weberi ) . The specimen is
attributed to "Ja^son" in the British Museum Catalogue.

Fig. 19. Ichthyophis javanicus sp. nov. Type. Ventral view
of caudal region; enlarged.

Ichthyophis malabarensis * sp. nov.

Type: Brit. Mus. Nat. Hist., No. 94.3.15.3, Maduvangard, Travan-
core, India; Ferguson, Collector.

Diagnosis: The largest oriental species known, reaching a length
of about half a meter. Tail proportionately long, its length in total
length 23.5 times; body width in total length about 27 times; trans-
verse primary and secondary folds, 360, 14 on tail; vertebrae. 111;
splenial tooth series, 10-10; other teeth numerous; tentacular open-

* Malabar -f- ensis, place locality of.

On the Caeolian and Related Species

Fig. 20. Ichthyophis maUibarensis sp. nov. Type. B. M. N. H. No. 94.3.15.3.
Maduvangard, Travancore, India. Actual length, 494 mm.

The University Science Bulletin

Fig. 21. Ichthyophis malabarensis sp. nov. Type. Radiograph
showing 111 vertebrae.

On the Caecilian and Related Species 83

ing close to edge of lip, nearer eye than to nostril; scales throughout
body in four to five rows in each fold.

Description of type: Head broadly oval; the distance between
the eyes (11 mm.) greater than length of snout in front of eyes
(8.4 mm.); tentacle situated close to lip, closer to eye (2.6 mm.)
than to nostril (5 mm.); distance between eye and nostril, 6.7 mm.;
width of head at first transverse groove, 14 mm., the head length,
21 mm.; tip of snout to first groove, 19 mm.; to second, 26 mm.; to
third, 32.3 mm.; two transverse folds appear anterior to the third
nuchal groove that are incomplete; 360 primary and secondary folds
encircle body forming on venter obtuse angles directed backwards,
meeting on the mid-line except on posterior part of body where
they pass straight across venter; on dorsal surface they are con-
tinuous, but anteriorly on the median line they tend to form a very
obtuse angle, directed forward.

Scales are present throughout the body, those on anterior folds
being small, sparse; posteriorly the scales increase in size and
number of rows in each fold. Posteriorly the scales may measure
2.5 to 3 mm. in diameter. They tend to form four or five imbri-
cating rows, the scales of one row overlapping laterally those of
the same row. The series of one fold tend to overlie partially those
of the following fold; vent longitudinal, interrupting six folds,
which tend to bend forward on venter; no glands visible on re-
gion of vent; tail pointed, somewhat laterally compressed, with 14
folds, the terminal portion without folds.

Teeth: maxillary-premaxillary, 28-30; vomeropalatine, 30-28;
mandibular, 28-28; splenial, 10-10; the maxillary-premaxillary teeth
are larger than the vomeropalatine; the mandibular considerably
larger than the maxillaries; the splenials distinctly smaller than the
vomeropalatines.

Coloration in preservative: Dark brown dorsally, the anterior
part of each fold darker than the posterior; laterally the color is
lighter and becomes nearly cream on underside of body. Head
above and on side, and anterior part of body somewhat mottled; a
transparent ring around eye, showing the bony border of the orbit
continuous; a cream spot on nostril; a small inconspicuous cream
mark around vent.

Measurements in mm.: Total length, 494; tail, 21; body width
(average), 18; head length, 20.2; head width, 14.5.

Remarks: It would appear that this species is rare in its range
or that it burrows to a considerable distance below the surface of

The University Science Bulletin

the earth. A creature so conspiciioiis would otherwise be better
known.

Despite the fact that it is the largest caecilian reported from Asia
it has a relatively low number of vertebrae (111). The number
of folds likewise is low when compared with certain other Indian
forms treated here in this paper. The number of folds on the tail
is 14, likewise a lower number than occurs in certain other Indian
forms.

Fig. 22. Ichthyophis malaharensis sp. nov. Type. Ventral
view of caudal region. Enlarged.

The specimen is a female containing many large eggs (5-6 mm.
in diameter), the number estimated to be more than 60.

The figure of Ichthyophis ^ijutinosus given in Wiedersheim (Die
Anatomic der Gymnophionen. Taf. B, fig. 28) shows the orbit in-
complete and continuous with the tentacular depression. It is
probably impossible to determine the source of Wiedersheim's ma-
terial. In this species the orbit is complete.* The mottled ap-
pearance, darker brownish marks on the anterior third of the body,
is I believe a normal coloration and not due to preservation. The
marks are more clearly evident when the specimen is submerged in
a clear liquid.

Ichthyophis youngorum sp. nov.

Type: EHI-HMS No. 35946, Doi Suthep (Sutep), Chiang Mai,
Thailand, at approximately 1200 m. in elevation; collected by Ed-
ward H. Taylor, July 12, 1957.

* The orbit is also complete in a specimen of 7. singaporensis as well as perhaps in
other species now placed in this genus. One may question whether these are congeneric with
Wiedersheim's specimen.

On the Caecilian and Related Species

Fig. 23. Ichthyophis youngorum sp. nov. Type. E. H. Taylor No. 35946,
Doi Suthep, Chiang Mai, Thailand. 3900 ft. elev. Actual length, 210 mm.

86 The University Science Bulletin

Paratypes: EHI-HMS Nos. 35944 adult; Nos. 35932-35941 larvae
all topotypes, July 12-15, 1957, same collector.

Diagnosis: A medium-sized species having large larvae (to 240
mm. ) ; primary and secondary folds, 314-326; body width in length
16-17.5 times; splenial teeth, in adults to 12-12; largest adult seen,
220 mm. Scales absent in at least anterior third of body; reduced
to a single overlapping row in each fold where present; vertebrae
106-107.

Description of type: A species of medium size, the body width
in length 17.5 times; width of head, 9 mm., its length, 12 mm.;
distance between eyes, 4.9 mm.; length of snout, 4.3 mm.; tentacle
small, cone-shaped, the opening lunate, closer to eye (1.3 mm.)
than to nostril (2.6 mm.).

First nuchal groove dimly visible on throat. Second groove
visible on throat but reaching the level of the mouth on side; third
groove barely indicated, not strong even laterally; a longitudinal
groove on chin extending on to throat. Snout-tip to third groove,
measured laterally, 16.5 mm.

The primary and secondary folds total 324 but many fail to reach
venter. The count on side is 304, on venter 280; four folds inter-
rupted by longitudinal vent; six folds on tail; a pair of dorsolateral
ridges indicated; transverse folds meet on venter at a broad angle
except posteriorly where they pass straight across venter. Scales
absent anteriorly, present in latter two thirds of body; when they
begin they are small, much wider than long; posteriorly they are
larger, with a single overlapping row in each fold; vertebrae 107.

Teeth: maxillary-premaxillary, 21-22; vomeropalatine, 21-22;
mandibular, 20-20; splenial, 12-12.

Color in life: General color above, violet to lavender; the venter
lavender; the grooves (partly due to glands in the skin) appear
light gray; chin blackish brown; head darker, nearly uniform
plumbeous; a gray-white spot in front of eye more or less con-
nected with a gray spot about tentacle; a slightly distinct light area
about nostril; the edges of the vent are gray flesh with two small
glands.

Measurements in mm.: Total length, 210; tail, 5.2; width of body,
12; width in length, 17.5 times; head length, 12; head width, 9;
tail length in total length, about 40.4 times.

Variation: Another specimen, an adult topotypic paratype No.
35944 is almost an exact counterpart of the preceding specimen.
It measures 220 mm. in length and the body width is contained in

On the Caecilian and Related Species

Fig. 24. Ichthyophis youngorum sp. nov. Type. E. H. T. No. 35946;
ventral view. Actual length, 210 mm.

The University Science Bulletin

the length 18 times. The count of the transverse folds on the dor-
sum is 328, the lateral count, 292, the ventral count, 285; vertebrae
106.

The following table of data gives the size and other variations
in a series of larva taken within a few meters of the adults. All are

Fig. 25. Ichthyophis youngorum sp. nov. Paratype, larva; E. H. T. No.
35940. Topotype. Lett figure, dorsal view; right figure, ventral view. Actual
length, 217 mm.

On the Caecilian and Related Species

Fig. 26. Ichthyophis youngorum sp. nov. Larval paratype E. H. T. No.
35940. Radiograph showing 107 vertebrae. Actual length 217 mm.

The University Science Bulletin

from a mountain rivnlet, the larvae having been taken from imdcr
rocks, trash or gravel in the stream bed.

The larvae have a single gill opening (spiracle) on each side
flanked by two small fleshy lobes. The nenromast system is pres-
ent on the head of the yonngest larvae.

The eye is represented in the larvae by a gray spot and only rarely
can the ontlines of the pnpil and iris be discerned. In yoimg larvae
the tentacular opening has not appeared; in older ones a depres-
sion may be found in the eye spot, and in the still older ones there
is a tentacular opening, usually curved, in the gray eye-spot. The
caudal fin is low and even in the youngest it can scarcely be traced

Fig. 27. Ichtliyopliis youngoiuin sp. nov. Larval paratype. E. H. T. No.
35949; topotype. Dorsal, lateral, and ventral views, showing the nenromast
system of young larvae.

below the tail. In the older larvae the fin has completely disap-
peared. The young larvae are lighter in color than adults, while
the older larvae may be a shade of ultramarine in color.

The largest larva 240 mm. has 106 vertebrae. Five larvae were
X-rayed and the vertebrae were 106 or 107.

Specimens were kept alive for a time in shallow water. They
were constantly moving when in the light. A slight noise could
be heard when they were taken from the water, presumably caused
by the water moving through the spiracle. No eggs were found.

This species is named for the family of Oliver Gordon Young
of Chiang Mai, Thailand. Mr. Young has been untiring in his
efforts to assist my \\ ork.

On the Caecilian and Related Species

Table

2. — Data

on Ichthyophis youngorum sp.

nov.

Width

NUMBEB

Length

Width

length

Number

Length

Width

in
length

35930...

35928. . . .

119

6.3

18.8

35927...

5.2

35934....

127

7.5

16.9

35947...

5.2

35931....

142

8.7

16.4

35949...

35932....

147

9.2

35929...

35935....

162

35933...

5.2

16.4

35937 ....

184

10.2

35948. . .

E>H

-4-^

o
o

— CO

3
O
u

a
o

CO — ;

33 o3

35936

194

17.6

325

280

14-14

18-18

12-12

35941

195

16.2

317

276

13-14

18-19

19-19

10-10

35942

213

12.5

314

302

16-16

16-18

19-19

10-10

35939

214
214

12.5
12

17.1
18

324
320

303

284

35938

14-14

16-17

19-i9

9-9

35940

217

12.2

17.8

313

304

16-16

18-18

19-19

10-10

35943

217

12.2

17.7

316

283

16-16

.19-19

19-20

10-10

35945

240

310

278

14-16

19-19

20-20

9-8

Ichthyophis sikkimensis * sp. nov.

Type: California Academy of Sciences, No. 64216, Darjeeling,
India.

Paratypes: British Museum No. 87.11.2.28, Darjeeling, Bengal,
India. Museum Comparative Zoology, No. 2685, Rungeet Valley,
British Sikkim, Tom Barbour collector; Berlin Museum, No. 2574,
Sikkim.

Diagnosis: A medium-sized species, characterized by 106-108
vertebrae; primary and secondary transverse folds 276-292; series
of splenial teeth (9-9 or 10-10); tail very short, contained approxi-
mately 50 times in total length, bearing five or six folds from front
of vent; tentacle near lip, closer to eye than to nostril. Scales
sparse or absent in anterior half of body; two to four rows in each
fold posteriorly.

Sikkiiim + ensis (Latin) = place or country.

The UNiNERSiri' Science Bulletin

Description of type: A medium-sized species, largest known
specimen, 276 mm. in total length; head slender, its width at first
annular groove (10 mm.), less than length of head (11 mm.); eye
closer to lip than nostril; tentacle very close to lip, nearer to eye
(1.7 mm.) than to nostril (2.9 mm.); distance between eyes (6.1
mm.) greater than length of snout (5.2 mm.).

First annular groove distinct laterally, dim on throat, not com-
pletely crossing head; second groove crosses throat passing up on
sides of head but a short distance; third groove distinct laterally
not completely crossing neck either above or below; first two or

Fig. 28. Ichthyophis sikkimensis sp. nov. Faratype.
3574. Left figure, dorsal view; right figure, ventral view.
270 mm.

Berlin Mus. No
Actual length.

On the Caecilian and Related Species

Fig. 29. Ichthyophis sikkimensis sp. nov. Paratype. B. M. N. H. No.
87.11.2.28. Radiograph sKowing 108 vertebrae.

three transverse folds not meeting below; primary and secondary
folds forming a slight median ventral angle, the grooves not meet-
ing on venter except posteriorly; (ordinarily, unless the specimen
is somewhat dessicated the meeting of the folds on venter can be
seen only dimly or not at all); total primary and secondary folds,
284, four of the posterior ones interrupted by vent; six folds on
tail counting from front edge of vent; tail-tip rather conical; a
slight longitudinal groove or depression on chin and throat.

94 The University Science Bulletin

Teeth: maxillary-premaxillary, 23-23; vomeropalatine, 21-21;
mandibular, 20-21; splenial, 10-10.

A pair of small pimples ( glands ) on each side of vent.

Color: The specimen has been long preserved. The color is
dark brown ( a little lighter on the ventral surfaces ) . The under-
side of chin still lighter, while the grooves on chin and neck are
marked dimly with cream; a more or less distinct cream tip on tail;
a narrow cream ring about eye; a small cream spot about nostril
and tentacle; the lips and tip of snout very light tan to cream.

Measurements in mm.: Total length, 276; tail, 5.5; width of body,
12.2; head width, 10; head length, 11.

Variation: No. 2685 has the pharyngeal region considerably
thickened and widened. The eye is milky white, the pupil not
visible. The tongue seemingly is not completely developed. It is
very short covering the splenial teeth, and its posterior limit is a

Fig. 30. Idithyophis sikkimensis sp. nov. Type. Ventral view
of caudal region. Enlarged.

ridge curving forward. This specimen agrees reasonably well with
the others in tooth counts, vertebrae and transverse folds. It is,
judging from the tongue, a recently transformed specimen that has
not attained all the adult characters.

No. 3574, the other Sikkim specimen, is very light, almost white,
on the venter and nearly white on chin. I cannot be certain that
this specimen has not been faded somewhat by light.

The two Darjeeling specimens vary but little from each other.

Nothing is known of the exact habitats except that No. 2685
comes from the Rungeet Valley.

On the Caecilian and Related Species

Data on type and paratype of Ichthyophis sikkimensis

Ichthtjophis s^umatramis* sp. nov.

Type: U. S. Natural Museum No. 70672, Kapahiang, Sumatra;
H. C. Kellars, collector.

Parotypes: U. S. N. M. Nos. 70667, 70669, Kaba Wetan, Sumatra;
70670, Kapahiang, Sumatra; all collected by H. C. Kellars.

Dmgnosis: A medium-sized species characterized by an elon-
gate series of splenial teeth, in adults 10-10 to 13-13; vertebrae,
110-112; transverse folds, females, 328-329; males, 315-318; tail in
total length in larger adults about 54-57 times; body width in total
length, average about 28 times; a semicircular row of small cream
glandules partly surrounding eye; largest specimen, 285 mm. in
length.

Description of type: Eye distinct, the lens appearing white, sur-
rounded by a black iris; a small whitish spot in front of eye; a
continuous semicircular row of tiny glandules run up from behind
eye, then forward, terminating in the light mark in front of eye,
(right eye abnormal); tentacle somewhat rounded at tip rather
than conical, the tentacular opening lunate, curving above tentacle,
opening close to lip at a point closer to eye (1.7 mm.) than to nos-
tril (2.8 mm.); distance between eyes measured on a curve, 5.5
mm.; length of snout, 4.4 mm.; distance between nostril and eye,
4 mm.; tip of snout to first nuchal groove, 11.3 mm.; to second,
14.6; to third, 18.5; first groove not visible above, moderately dis-
tinct ventrally; second very distinct ventrally, slightly so on side
of neck; third only indicated laterally.

* Sumatra -|- anus (Latin) = place, location, country.

The University Science Bulletin

Fig. 31. Ichthyophis sumatranus sp. nov. Type. U. S. N. M. No. 70672,
Kapahiang, Sumatra. Actual length, 273 mm.

On the Caecilian and Related Species 97

The first four or five transverse folds do not meet on venter;
other folds cross back almost directly, curve back ventrally and
form a median ventral angle, three to four millimeters behind
the point where they crossed above; in the posterior part, the folds
pass around body in about the same plane without forming an angle;
vent interrupts five or six folds; total transverse folds, dorsal count,
315; 7 folds on tail; the transverse grooves rather dim (the speci-
men has been dehydrated slightly ) and in places cannot be clearly
discerned. They are distinct on sides and in the posterior part of
body. The specimen is a male.

Small scales present anteriorly but sparse on dorsal and ventral
parts of folds; more posteriorly they increase in size and number
until there are three to five transverse imbricating rows passing
around the body in each fold, partially overlapping the series of
scales of the following fold, but separated from them by a row of
recumbent glands lying nearly longitudinally.

Teeth: maxillary-premaxillary, 22-21; vomeropalatine, 22-21;
mandibular, 20-19; splenial, 13-13. Vertebrae 112.

Color in preservative: Above dark brown, the grooves slightly
lighter; ventral coloration a lighter shade of brown; a cream mark
at vent, and at tip of tail; a tiny spot of white in front of eye and
a fine semicircular row of cream-colored glandules partly sur-
rounding eye; a cream spot at tentacle and nostril; edge of lips
cream.

Measuremenis in mm.: Total length, 273; tail, 5; body width,
10; head length, 10.5; head width, 8.6.

Remarks: There are three paratypes, one a topotype from Kapa-
hiang, a female, and two from Kaba Wetan.

No. 70667 the smallest specimen has had the internal organs re-
moved. Where sex has been determined females have the higher
count of transverse folds. The series is too small to postulate that
this is generally true. This specimen has the eye-area milky white,
the eye being distinguished with difficulty. In consequence the
semicircular series of glandules cannot be seen. The specimen is
violet-plumbeous in color. These seeming differences from the
type may be due to age and method of preservation.

No. 70672 has the head somewhat lighter than body. The tip
of the snout is somewhat cream-colored. This specimen shows a
lighter line along the grooves, a character not or less evident in the
smaller specimens.

The following table presents characters of these specimens.

4—8920

The University Science Bulletin

Table

of data fi

"■oin paratypcs

of Ichthyophis sumatranus

Number

Length

Tail

Bodv

width

Width

in
lengt h

Head
width

Head
length

70669 . .

285
273

5
5

9.5

27 3

8.6

11.5

70672

10.5

70667

205

3.5

25.5

7.2

10.1

70670

176

3.2

NUMBEK

Verte-
brae

Bodv
folds

Tail
folds

Max-
pre-
max.

Vomero-
palatine

Man-
dibular

Splenial

70669

112

329

21-21

23-22

12-13

70672

112

315

22-21

20-19

13-13

70667

110

328

22-22

19-19

20-20

12-13

70670

110

318

20-21

24-23

18-19

10-10

Ichthyophis acuminatus* sp. nov.

Type: American Museum of Natural History, No. 20875, Me
Wang Valley, Thailand, Malcolm Smith, collector.

Paratypcs: British Museum (Natural History), No. 1921.4.1.338,
Me Wang, N. Thailand (field No. M. S. 3135); Malcolm Smith
field Nos. 5656 Muang Liep, Thailand, and 3185, 3187 Pa Meang,
Me Wang, Thailand.

Diagnosis: Large ( about 300 mm. ) ; head rather acuminate; eyes
visible, very small; tentacle near lip, twice as close to eye as to nos-
tril; splenial teeth, in transformed specimens, 15-15 to 22-22 in old
adults; tail short, without cream spot about vent. Scales present in
posterior part of body, wanting or greatly reduced anteriorly; body
width in body length (in adults) about 20 times; transverse body
folds on dorsum, 315-330, on venter, 297-320; vertebrae, 109-110.
Larvae transform at a length of about 205 mm. Tail length in total
length, approximately 43 times.

Description of type: Body thick, short, with dorsolateral ridges
evident along sides; head acuminate, and, seen from above, form-
ing a triangle; eye visible, minute ( .7 mm. ) ; tentacle close to lip.

acuminatus (Latin) ;r pointed, referring to the snout.

On the Caecilian and Related Species

Fig. 32. Iciithtjophis acunundtus sp. nov. T\ pc. A. M. X. H. No. 20875.
Mc \\ang \'allty, Thailand. Actual length, 295 mm.

100 The University Science Bulletin

minute, conical, the opening somewhat horseshoe-shaped, closer to
eye (1.7 mm.) than to nostril (3.8 mm.); width between eyes, 7.8
mm. (measured on curve); length of snout, 5.8 mm.; tip of snout
extending beyond mouth, 1 mm.; width of head, 11.6 mm.; length
of head, 14 mm.; first annular groove well defined on sides of head
and under chin; second groove ventral, ascending on side of head
slightly, separated from preceding groove by a distance of 4 mm.;
third groove scarcely indicated; first primary fold does not cross
throat; all subsequent primaries and secondaries practically com-
plete ventrally; the anterior folds are dim or incomplete dorsally on
anterior fifth or sixth of body; the folds slightly angled on venter,
while elsewhere they pass nearly straight across body; 315 primary
and secondary folds, counted dorsally; 303 counted on venter; six
confined to tail. Tail length in total length approximately 43
times.

Scales present in posterior two thirds of body, one or two rows
to each fold; if present anteriorly scales much reduced, not extend-
ing on to venter and usually the scales in each transverse row are
not contiguous; vent longitudinal, interrupting six folds; tip of tail
pointed, flat on ventral surface, somewhat compressed laterally.

Teeth: maxillary-premaxillary series, 24-25; vomeropalatine, 26-
27; mandibular, 2.5-24; splenial, 22-22 much smaller than other
teeth. Tongue rather pointed, not covering the large series of
splenial teeth. Vertebrae, 110.

Fig. 33. Ichthyophis acuminatus sp. nov. Type. Ventral view of

caudal region. Enlarged

On the Caecilian and Related Species 101

Color in preservative: Somewhat violet-lavender, nearly uniform
above, tending to be slightly lighter on throat and chin; area about
vent somewhat lighter hut not cream; area about tentacular open-
ing cream; lips light flesh to cream.

Measurements in mm.: Total length, 295; tail, 7 (from front of
vent); head width, 11.6; head length, 14; body width, 14.6; body
width in length, 20.2 times.

Variation: There are two other transformed specimens from the
Me Wang Valley in northern Thailand and a small series of larvae
that most probably belong to this species. The larvae are listed
with the adults in the following table, showing comparative meas-
urements. The most significant variation (in teeth and folds) are
given. The difference in count on dorsum and on venter is such
that both counts are given. A lateral count is not likely to be
either higher or lower than these counts. The counts of teeth and
folds in the younger larvae are not trustworthy, hence are not
recorded in the table.

Ichthyophis nigrofavus * sp. nov.

Type: U. S. N. M. No. 129462 "within 20 miles of Kuala Lumpur,"
Selangor, Malaya; Traub and Tipton collectors.

Diagnosis: A large species (425 mm.) the width contained in
length about 26.5 times; body folds, dorsal count, 416 (ventral 399);
folds on tail 7-8; vertebrae 123; maxillary teeth, 31-32; vomeropala-
tine 32-33; mandibular, 28-29; splenial, 0-0. Black above and below,
a yellow lateral stripe beginning under eye narrow at first, widening
posterior to pharynx; eye, tentacle and nostril with gray-white rings
or spots ( not cream ) .

Description of type: Head oval anteriorly, the sides nearly par-
allel, its width at first nuchal groove 10.7; head length, 12.8; eyes
distinct, the lens white surrounded by a very narrow line of black
(iris); these surrounded by a complete ring of gray-white; the
distance between eyes, measured on the curve ( 8.5 mm. ) , greater
than snout length (6 mm.); eye and nostril equidistant from lip,
the distance between them, 4.9 mm.; tentacle conical the opening
lunate, situated closer to eye (1.5 mm.) than to the nostril (3.8
mm.); tip of snout extends beyond nostril 1 mm.; tip of snout to
first nuchal groove, 14 mm.; to second, 18.2 mm.; to third, 22.2 mm.

Dorsal count of primary and secondary folds (they cannot be
distinguished), 416; counted on venter, 399; folds on tail, 8; scales

* nigro (Latin) = black -)- flavus (Latin) ^ yellow

102

The University Science Bulletin

CO
1—i

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1-H
»-H

re >0 C<1 ■*
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1 1 1 1
<N IC O »0
C^ (N <N C^

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1 1 1 1
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On the Caecilian and Related Species 103

are absent or if present minute on anterior sixth of length; more
posteriorly they increase in number and size and in the posterior
part of the body there are four or five transverse overlapping im-
bricating rows in each fold. There are 123 vertebrae. First nuchal
groove surrounds head; second not visible above; third distinct
laterally but can be traced across throat.

There are no splenial teeth, but in the other series the number
of teeth exceed any other fonn described from southeastern Asia
and the archipelagoes.

Teeth: maxillary-premaxillary, 31-32; vomeropalatine, 32-33;
mandibular, 28-29; splenial, 0-0.

There are two small glands present on each side of the vent in
its anterior part; the vent interrupts three folds; the length of the
tail in total length, approximately 70 times.

Color in fixative: Dorsum and venter black with a cream lateral
stripe, the edges of which are very irregular. Anteriorly it begins
under eye, passes back to neck where it widens and continues to
level of vent narrowing posteriorly; a small cream spot at vent, at
tentacle and nostril; ring about eye gray-white; chin and throat
deep black. There are many minute black Hecks on the cream
stripe.

Measurements in mm.: Total length, 425; tail, 6; width of body,
16; width of head, 10.7; length of head, 14.

Remarks: The specimen has been somewhat dehydrated and in
such places the black has become olive, and the ventral black
shows a whitish surface.

Another specimen U. S. N. M. No. 129463 bears a similar label
l)nt this is unquestionably a different species. It diflFers among
other characters in having approximately 100 less folds and there
is a series of splenial teeth present.

The characters of the black color, the large number of vertebrae
(123), the large number of folds, and absent splenials set this spe-
cies apart from other Asiatic forms with a lateral stripe.

Ichthyoj)Jiis paucisulcus * sp. no\ .

Type: U. S. National Museum No. 103565; from Siantar, Sumatra;
National Geographical-Smithsonian Institution Expedition, coll.

Diagnosis: A short, broad species the width in length about 17
times; transverse folds on body, 259 dorsal count, 263 lateral count,
folds on tail, 5-6; tail length in total length, approximately 54 times;

* pauci ( Latin ) = few + sulcus ( Latin ) ^ groove.

104

The University Science Bulletin

splenial teeth 14-14; choanae oval (not angular); grooves not meet-
ing on venter except in posterior part of body; folds forming a very
obtuse angle on median ventral line except posteriorly where the
grooves and folds go straight across venter. A narrow cream lateral
stripe, not broken on neck.

Description of type: Head nearly as broad (11.2 mm.) as long
(12.6 mm.); eye-spot elevated, circular, showing a darker center
and a ring of cream; tentacular opening lunate, situated near lip,
distinctly closer to eye (1.6 mm.) than to nostril (3.7 mm.); dis-
tance between eyes (7.4 mm. measured on curve) much greater

Fig. 34. Ichthyophis paucisulcus sp. nov. Type. U. S. N. M. No. 103565,
Siantar, Sumatra. Actual length, 256 mm.

On the Caeciliax and Related Species 105

than snout length (5.2 mm.); nostril 1.2 mm. distant from lip; eye,
1.8 mm. from same; eye from nostril, 4.7 mm.

Snout to first nuchal groove, 13 mm., to second, 16.8 mm., to third,
20.2 mm. (the measurements made on side of head); first groove
distinct, passing entirely around head; second distinct below and
on sides to level of mouth; third distinct laterally, absent on throat,
dimly indicated above; folds forming an obtuse angle on venter
except posteriorly where they are straight; grooves cannot be traced
across venter in anterior half of body; folds, dorsal count, 259;
lateral count, 266. A few of the folds split on the side; vent longi-
tudinal, interrupting four folds. A pair of slight swellings on each
side of vent suggest the presence of special glands; 5 or 6 folds on
tail from front of vent; small scales are present on dorsum on an-
terior part of body but do not pass to ventral side; posteriorly the
scales increase in number of rows in each fold and they are con-
tinuous around body, as many as four or five rows present poste-
riorly.

Teeth: maxillary-premaxillary series, 28-28; vomeropalatine, 26-
27; mandibular, 28-28 the posterior teeth very small; splenial, 14-14.
The tongue (injured) has a triangular thickened anterior part that
probably covers the splenial tooth series; the remainder of the
tongue is thinner and striated.

Color in preservative: Above lavender-slate; a longitudinal cream
stripe beginning as a narrow line below eye, passes back laterally
and is interrupted at the third costal groove; the stripe widens and
terminates laterally at level of vent, the edges of stripe not discrete;
entire venter with rather indistinct very numerous markings of
lavender and tan, not strongly contrasting, placed transversely; a
well-defined cream spot surrounds vent; a cream spot at tentacle;
tip of snout light; the lower lips cream, the jaws grayish below with
a grayish spot on chin. The stripe does not bifurcate at angle of
the mouth.

Measurements in mm.: Total length, 256; tail, 4.7; body width
average about, 15; width in length about 17 times; head width, 11.2;
head length, 12.6; tail length in total length approximately 55 times.

Remarks: A specimen obtained by W. L. Abbot on North Pagi
Island in the southeastern part of the Mentawai group may belong
subspecifically with the preceding species. It differs in having teeth
and bones green, a condition most probably caused by some acci-
dent of preservation. Tlie characters of this specimen show some
significant variation. It is slenderer proportionally, the head longer,

106

The University Science Bulletin

Fig. 35. Ichthyophis paucisulcus sp. nov. A specimen U. S. N. M. No.
31701 from Pagi I, Mentawai Group, Sumatra, referred to this species. Actual
length, 210 mm.

On the Caecilian and Related Species 107

slightly fewer body folds (9). The vertebrae number 104. The
lateral light stripe is narrow and indistinct.

Total length, 201; tail, 4; tail in length, 50 times; body width,
average, 10; width in length, 20 times; head width, 9; head length,
12.4; tip of snout to first groove, 9.6; to second groove, 12.6; to third,
15.2; total folds on body, 250; folds on tail, 8. Teeth: Maxillary-
premaxillary series, 22-20; vomeropalatine, 22-22; mandibular, 22-
23; splenial, 10-10. Tongue somewhat pointed, not covering splenial
teeth.

Idithyophis stipachaii sp. nov.

Type: EHT-HMS No. 35498, 10 km. west Nakon Si Thamarrat,
Nakon Si Thamarrat province, taken Apr. 30, 1958 by E. H. Taylor.

Parahjpes: EHT-HMS Nos. 35497, 35499. Topotypes same date
and collector. No. 34677 young, transformed 18 km. N. E. Betong,
Yala, Nos. 35781-82 larvae and one transformed young. No. 35780
Kao Chao Forest Station near Trang, Trang Prov., Nos. 35594-96
larvae and No. 35593 young transformed specimen, Rompibong Tin
Mine, Nakon Si Thammarat.

Diagnosis: A relatively slender, elongate species, the largest spec-
imen known, 306 mm., with a lateral cream stripe broken on neck;
numerous scattered cream spots on dorsum and venter; width in
length, 30.6; tail in length 102 times; mandibular teeth reduced in
number, the splenials 18-18 becoming more prominent; folds 322;
larvae transform at small size (before a length of 125 mm. is
reached); tentacle nearer to eye than to nostril.

Description of type: Head rather flattened, its greatest elevation,
3.2 mm.; width of head (9.2 mm.) less than the length (13 mm.);
distance between eyes (6.2 mm.) greater than length of snout in
front of eyes (5 mm.); tentacle conical, the opening somewhat
lunate, closer to eye (1.8 mm.) than to nostril, 3.5 mm.; distance
between eye and nostril, 4.8 mm.; tip of snout to first nuchal groove,
12.2 mm.; to second, 16.4 mm.; to third, 20 mm.

Primary and secondary folds, 322, with 4-5 on tail; the grooves
failing to meet dorsally or ventrally anteriorly; scales absent in an-
teriormost folds, appearing first dorsally where they are in a single
row in each fold, and about one-half millimeter in diameter; pos-
teriorly they are both dorsal and ventral, sunounding body, in from
three to five rows in each fold, overlapping transversely.

Teeth: maxillary-premaxillary, 31-32; vomeropalatine, 27-27;
mandibular, 8-6; splenial, 18-18. Tongue oval, rather narrowed
anteriorly, not covering, in fact not reaching, the splenial teeth.

108

The University Science Bulletin

Fig. 36. Ichthyophis supachaii sp. nov. Type. E. H. T. No. 35498, 10 km.
W Nakon Si Thammarat, Thailand. Actual length, 306 mm.

On the Caecilian and Related Species

109

Color in life: Above dark plumbeous lavender; gray slate below;
a spot of cream near mouth angle; another spot on side of pharynx;
a cream lateral stripe begins at the third nuchal groove and con-
tinues to a point somewhat in advance of the vent; numerous flecks
and spots of cream on venter, a few present on dorsum. Eye with
a pearly lens surrounded by a fine ring of black, and a narrow, in-
complete ring of cream about eye, a cream spot about tentacle and
one about vent; extreme tip of tail light; a small median cream spot
on occiput.

Measurements in mm.: Total length, 306; tail length, 3; width
of body, approximately, 10; width in length, 30.6 times; tail in
length, 102 times; width of head, 9.2; length of head, 13.

Fig. 37. Ichthyaphis supachaii sp. nov. Paratype larva, E. H. T. No. 35781,
near Trang, Trang Prov., Thaijand. Actual length, 97 mm.

Remarks: Two small specimens, one recently transformed, were
taken with the type in clay soil on the bank of a small stream. It
would appear that the species completes its transformation when
it is less than 125 mm. in length.

Measurements of EHT-HMS No. 35499 topotype: This is the
smallest of the two topotypes and has the following characters:
Total length is 125 mm.; the tail, 2.8 mm. (a portion of the caudal
fin is still evident in a compressed ridge that extends along the dor-
sal surface of the tail reaching the tail tip ) . The width of the body
is about 5 mm., the width of head, 4 mm., the length, 7 mm.

110 The University Science Bulletin

The transverse folds number 313, seven being confined to the
tail. The folds meet in an obtuse angle ventrally as they do in the
type.

The teeth foreshadow the condition obtaining in the adult but
all the series are reduced. I count only five maxillary-premaxillary
teeth on each side. In the vomerine series there are seven on each
side. In the mandibular series I find a pair of very tiny teeth near
the symphysis, then three widely spaced enlarged teeth. In the
posterior part of jaw there is a single tooth visible on one side, two
on the other. The splenial series has two enlarged anterior teeth
(as large as the mandibular), followed by three symmetrically
spaced teeth on each side. The tongue lies completely behind the
splenial row, and is poorly developed at this stage.

The young larval specimens do not show the white spotting on
dorsum or venter. There is a small cream mark running from the
eye (which is distinct) to the tentacle. The gill opening is single.

The characteristics of the mandibular teeth distinguishes this
species from all others here recognized.

The species is named to honor Prof. Supachai Vanijadhana, Sec-
retary General of Chulalongkorn University, whose interest in the
faunas of Thailand made possible my journeys in that country.

Ichthijophis kohtaoensis sp. nov.

Type: U. S. National Museum No. 72293, Koh Tao Island, west
side, Gulf of Siam, Malcolm Smith collector, (field number 2932).

Paratype: U. S. National Museum No. 76138, topotype, same col-
lector.

Diagnosis: A form with lateral stripes; a large series of splenial
teeth (17-18); number of transverse folds, lateral count, 362-366;
tentacle small, opening near lip, closer to eye than to nostril; eye
very distinct; light stripe widens on side of head and bifurcates at
mouth, terminating anteriorly below eye; posteriorly, somewhat in
advance of level of vent; width in length about 24 times; known
length, 280 mm.

Description of type: Head rather narrow, the pharyngeal region
a little wider than head; width of head at first groove, 8.7 mm.;
length of head, 12.1; width between eyes, 5.8 mm.; length of snout,
4.4 mm.; tentacular opening curved, small, near edge of lip, closer
to eye (1.7 mm.) than to nostril (3 mm.). Eye distinct, the lens
light surrounded by a narrow black rim, the eye itself surrounded
by a narrow cream ring. First nuchal groove on neck, distinct on

On the Caecilian and Related Species

111

Fig. 38. Ichthyophis kohtiioensis sp. nov. Type. U. S. N. M. No. 72293,
Koh Tao Island, west side, Gulf of Siam. Actual length, 280 mm.

sides and below; second groove strong below, reaching above the
lateral light line; third groove can be traced around neck except on
the median ventral point; three dorsal folds lie between the second
and third nuchal grooves; the subsequent folds turn back somewhat
on venter and meet on midventral line at an angle; the grooves fail

112 The University Science Bulletin

to cross the ventral surface except posteriorly in front of vent where
the folds and grooves cross the venter in a straight line; longitudinal
vent interrupts about four folds. Total transverse folds (dorsolat-
eral count including the three dorsal ) 362 of which six are confined
to the tail; tail narrowing suddenly to a point, the subcaudal region
flattened.

Scales present anteriorly ( seemingly none on the three pharyngeal
folds); at first an occasional small scale is present; more posteriorly
scales increase in size and in number and two to three (perhaps
four) rows of imbricating scales varying in size, are present in each
fold the rows overlapping and extending around the body.

Teeth: maxillavy-premaxillary, 22-2-3; vomeropalatine, 22-23;
mandibular, 21-20; splenial, 17-18. Vomeropalatine teeth scarcely
extending through gums.

Color in preservative: Generally lavender brown, the grooves
lighter giving an effect of minute bands of light and dark; a broad
lateral stripe of cream from eye to a point a little in advance of
level of vent; a cream area about vent; venter same shade as dor-
sum; the lateral stripe widens behind mouth angle and bifurcates,
the lower branch short, quickly becoming brownish; a small cream
ring about eye; a cream area around tentacular opening, and one
around nostril; lips and tip of snout light brown to brownish cream.

Measurements of type and parattjpe in mm.: Snout to vent, 280;
192; tail, 4.2; 3.3; widtli of head, 8.7; 6.9; length of head, 12.1; 9.6;
width of body, 12.2; 8.2; width in length (times), 23; 23.4; tail
length in total length (times), 66.6; 60.

Remarks: The paratype has a pair of triangular yellow cream
spots on each side and slightly behind vent, of a shade lighter than
the spot about vent. There are 366 folds (dorsolateral count). The
teeth are: maxillary-premaxillary, 21-22; vomeropalatine, 16-17;
mandibular, 19-19; splenial, 13-14. These specimens are from Koh
Tao, a small island some 70 km. off the eastern shore of peninsular
Thailand.

They differ from the mainland form chiefly in having a larger
number of transverse folds on the body. The largest count made
for specimens from the mainland of Thailand nearest these islands
is 323 a difference of about 39 folds. Farther to the northeast in
specimens from Viet Nam my highest count was 344.

The island of Koh Tao would appear to have had a somewhat
different history from many of the islands in the Gulf of Siam. A
number of its forms of reptiles and amphibians differ so much from

On the Caecilian and Related Species 113

the mainland species that they have been recognized by species or
subspecies names. As yet the island is little explored. The species
is named from the island + ensis ( Latin ) place, country.

Ichthyophis beddomei Peters

Ichthyophis Beddomii Peters, Monats. K. Akad. Wiss. (Sitz. phys.-math.
Classe), Nov. 1879, p. 932, pi. fig. 4 (type locality Nilgherries, India).

Diagnosis: A species having 240 transverse folds; the tentacular
opening almost equidistant from eye and nostril; snout more pointed
than in glutinosa; splenial teeth well developed; length, 225.

Description: "Nur 240 Hautfalten. Die Tentacle-grube kaum
weiter von dem Nasloch als von dem Auge entfemt. Schnause
spitzer als bie der vorhergehenden Art. Wie bei der vorigen an
jeder Seite eine gelbe Langsbinde. Zvveite Reihe der Unterkiefer-
ziihne wohl entwickelt. Totallange 0,225; Kopf 0,011; Korperbreite
in der Mitte 0,010.

"Von den Nilgherries. Wir verdanken diese Art der Giite des
Hrn. Colonel Beddome. (M. B. No. 5545)."

It seems almost certain that this species is one distinct from
glutinosa differing in having a hundred less transverse folds than
the Linnean species.

This species was synonymized by Boulenger but an examination
of the material that he studied now remaining in the British Mu-
seum shows that he has confused two or more forms.

Ichtlnjophis tricolor Annandale

Ichthyophis glutinosa tricolor Annandale, Rec. Ind. Mus., vol. 3, 1909, p. 286
(type locality, Maddathorai, India); idem., vol. 9, pt. 4, no. 19, Aug. 1915,
pp. 346-347 ( Western Ghats, Cochin ) .

Diagnosis: A species having a white venter.

Description: Annandale describes the form very briefly as fol-
lows: "A specimen was taken at Maddathorai in a hollow tree. It
had the whole of the ventral surface pure white, and therefore dif-
ferent in appearance from the typical form. A careful comparison,
however with normal specimens, including a microscopic examina-
tion of the scales, reveals no other difference. I propose to call
the form with the white ventral surface, — var tricolor."

Remarks: Annandale in 1915 reported two more specimens as
follows: "Two specimens of this variety or local race were found
by Mr. F. H. Gravely on the eastern slopes of the Western Ghats
in Cochin in September last, the exact locality being Parambikulam
(alt. 1700-3200 feet). The specimens are considerably larger than

114 The Uni\^rsity Science Bulletin

the type, one of them being 280 mm. long. The yellow lateral band
on each side is separated from the white median ventral band by a
dark one, which is greyish in spirit. This dark band varies consid-
erably in breadth."

The striking difference in coloration strongly suggests that we are
dealing with a distinct species rather than a color variety since
there are at least three other species of the genus occurring in the
same general locality. The color pattern is normally very constant
in caecilians. Since Mr. Annandale did not report on folds, teeth,
\ertebrae, and other significant characters, complete description
must await re-examination of the type. Moreover we do not know
what species Annandale used for comparison.

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On the Caecilian and Related Species 119

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120 The University Science Bulletin

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THE UNIYEESITY OF KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 5

A Review of the Genus Paroxystoglossa^
( Hymenoptera : Halictidae )

J. S. Moure, C. M. F.^

Department of Entomology, University of Kansas, Lawrence and Secg^ao de
Zoologia, Universidade de Parana, Curitiba, Brasil

Since the genus Taroxijstoglossa was described (Moure, 1940),
several additional species have come to hand. All known species
occur in southeastern South America, from Brasil to Argentina.
The species are enumerated and described below.

Paroxystoglossa jocasta (Schrottky)

Oxystoglossa jocasta Schrottky, 1910, Rev. Mus. Paulista, vol. 8, p. 74.
Halictus mallotes Vachal, 1911, Misc. Ent. [Narbonne], vol. 19, p. 42, 43, 50.
Paroxystoglossa jocasta jocasta, Moure, 1940, Arq. Zool. [Sao Paulo], vol.

2, p. 60; Moure, 1943, Rev. Ent. [Rio de Janeiro], vol. 14, p. 477.
Paroxystoglossa jocasta curitybana Moure, 1940, Arq. Zool. [Sao Paulo],

vol. 2, p. 60.
Paroxystoglossa jocasta, Moure, 1950, Dusenia, vol. 1, p. 311.

The type locality of this species is Campos do Jordao, 1600 meters
altitude, Sao Paulo, Brasil. Halictus mallotes was described by
Vachal from Parana and from "Bresil (Sello)," the latter specimen
being very probably from the state of Siio Paulo. The color variant
that I gave the name curitybana is from Curitiba, 900 meters alti-
tude, Parana, Brasil. The type of jocasta is in the Departmento de
Zoologia da Sccretaria de Agricultura, Sao Paulo; that of curitybana
is in my collection in Curitiba.

1. Contribution No. 981 from the Dcpartnit-nt of EntomoloRv, University of Kansas.

2. I wish to thank the Rockefeller Foundation (New York), the National Science
Foundation (Washington) and the Canipanha de Aperfeigoamento de Pessal de Nivel
Superior (Rio de Janeiro) for aid that made this study possible. Also. I wish to thank Dr.
Carlos Alberto Campos Seabra of Rio de Janeiro for the stimulus and generosity which he
is giving to studies of Brasilian bees, and Dr. Charles D. Michener of the University of Kan-
sas for help in preparation of this paper.

(121)

122 The University Science Bulletin

The pilosity and punctation are rather constant in this species,
in which, however, the metalHc coloration varies from hhie green
to golden green, and in many specimens the abdomen is more or less
strongly coppery red.

Distribution: Parana (Curitiba, Ponta Grossa, Palmeira, Co-
lombo, Campo Largo, Gnarapuava, Sfio Jose dos Pinhais), Sao
Paulo (Sao Paulo, Guarulhos, Campos do Jordao), and Minas
Gerais (Fazenda dos Campos, Passa-Quatro); all these localities
in Brasil.

Paroxystoglossa barbata sp. nov

Female: Color entirely metallic green, passing gradually to blue
on the marginal depressions of the tergites, their extreme margins
bleached; mandibles brown, reddish in the middle, with a green
spot basally; apical part of clypeus black with some red reflections;
scape brown, nonmetallic, flagellum paler brown; extremities of
tibiae and tarsi brown, nonmetallic on the posterior basitarsus,
sternites with median bands of greenish. Tegulae brown, the
anterior outer third translucent whitish, with small elongate green
spot on the anterior inner margin. Pilosity of wing membrane as
well as veins and pterostigma honey colored, vein R darker.

Pilosity white, well developed, especially on face, genal areas,
episterna, sides of propodeum and sternites; on the tibiae and tarsi,
as well as sides of rima of fifth tergite, very pale brown.

Punctation very dense and fine on the front and vertex, passing
to a little coarser and sparser on the lower paraocular areas; very
fine and dense on upper genal areas and much sparser below; on
clypeus and supraclypeal area coarser, although very sparse on
disc and finer at sides; on mesonotum a little less dense than on
frons or on scutellum, rather uniform, only a little sparser on an-
terior part; on mesepisterna dense above, considerably sparser be-
low anteriorly, finer and denser near metepisterna; metepisterna
roughly punctulate; sides of propodeum rough, postbasal angles
and posterior surface shining, microreticulate, piliferous punctures
fine. First two tergites with punctation much finer than on
mesonotum, rather dense, even on marginal depressions; terga
three to five a little more sparsely punctate, especially on discs.

Head distinctly broader than distance from clypeal margin to
anterior ocellus (143:105); upper interorbital distance greater
than lower, this a little less than eye length (90:82:85); interocellar
distance slightly less than ocellocular (24:26); scape longer than
half maximum width of face (65:53), surpassing vertex; frontal

Review of the Genus Paroxystoglossa 123

carina evident only below; malar area linear. Anterior part of
mesonotum with almost semicircnlar contours, little salient and
feebly depressed in middle; corners of pronotum forming very
obtuse angles with rounded apices; punctulation of metanotum
very evident. Basal area of propodeum shorter than metanotum,
with semilunar depression, posterior carina rather evident, forming
a broadly open V, surface roughened, with strong central carina
and vestigial basal radiating carinae.

Length 9 mm.; wing length including tegula 9.7 mm.; width of
head 2.4 mm., of abdomen 3.3 mm.

Male: Color as in female, with green spot on base of mandible,
scape metallic green and pedicel with greenish reflections; clypeus
wholly green; flagellum darker, especially basally; posterior basitar-
sus with green reflections on the outer face.

Pilosity white, even more developed than in female, especially on
genae and all the venter; on tibiae white, and on apical tergites with
some pallid hairs among brown ones.

Punctation as in female, but a little denser on frons; on supra-
clypeal area and clypeus intervals little broader than punctures,
dull reticulate on former; punctation stronger on episterna, well
defined and rather strong on sides of propodeum; on tergites a little
stronger but much finer than on mesonotum, even on sides of first
tergite.

Head distinctly longer than clypeocellar distance (145:105);
upper interorbital distance equal to eye length, considerably greater
than lower interorbital distance (85:85:70); interocellar distance
clearly less than ocellocular distance (23:26); scape less than half
as long as facial width but longer than two first flagellar segments
together; first flagellar segment a little shorter than second, which
is clearly shorter than first and pedicel together; frontal carina al-
most reduced to an interantennal tubercle; malar area linear; pre-
occipital carina absent. Anterior part of mesonotum a little pro-
duced, forming a semicircular prominence, the anterior median part
of which is slightly depressed. Femora and tibiae, especially of
posterior legs, a little swollen. Basal area of propodeum still more
prominently margined, semilunar, the carinae a little more evident
than in female. Metasoma with fourth and fifth sternites more
sclerotized medially and depressed forming a canal which narrows
toward apical margin of fifth sternite.

Length 9.5 mm.; wing length including tegula 6.7 mm.; head
width 2.4 mm.; abdominal width 2.8 mm.

124 The University Science Bulletin

Distribution: Tandil, 250 meters altitude (type locality), No-
vember, 1954 (F. H. Walz) and Lavallel, March 20, 1952 (M.
Senkute ) ; both localities in the Province of Buenos Aires, Argentina.

Types: Holotype, allotype, and a female paratype in the Snow
Entomological Museum, University of Kansas; two paratypes (male
and female ) in the collection of the author; one female paratype in
the Museo Nacional "Bernardino Rivadavia," Buenos Aires.

Paroxystoglossa transversa Moure

Paroxiisioglossa transversa Moure, 1943, Rev. Ent. [Rio de Janeiro], vol. 14, p.
477; Moure, 1944, Rev. Ent. [Rio de Janeiro], vol. 15, p. 275.

This species was described from Puerto Bertoni, Paraguay, and all
known specimens come from the Parana basin, several hundred
specimens being at hand from Nova Teutonia.

The variation in coloration in the females is extraordinary, as was
noted in 1944. Among specimens from Nova Teutonia at the Uni-
versity of Kansas are all color variations previously noted as well
as specimens which are entirely dark brown and some approaching
blue. Examples with the head and thorax green and abdomen red
are also relatively common.

In males the coloration is generally green, passing to gold, and
sometimes weakly coppery on the abdomen.

On the contrary the characters noted in the key are very constant,
although the size of the females varies considerably. This species
would seem to be very interesting for biological study in view of
the great size variation among females.

Distribution: Santa Catarina, Brasil (Nova Teutonia); Paraguay
(Puerto Bertoni and Hohenau).

Paroxystoglossa brachycera sp. nov.

Female: Color metallic green, sometimes with golden reflections
or even passing to blue green, tergal margins a little translucent;
mandibles brownish black, without metallic basal spot; anterior
third of clypeus dark with some purplish reflections; scape brownish
black, nonmetallic, flagellum rather dark; apices of tibiae and tarsi
brown, posterior basitarsi nonmetallic; sternites with median band
of metallic green, bleached. Tegulae dark brown with a green spot
anteromedially, translucent anterolaterally; veins, pterostigma, and
pilosity of wing veins pale honey colored, vein R a little darker.

Pilosity white, a little less abundant than in P. barbata and jocasta,
longer on genal areas, sides of thorax, and ventral sides of thorax
and abdomen than elsewhere, that of tibiae and tarsi and sides of
rima of fifth tergite very pale brown.

Re\t[ew of the Genus Paroxystoglossa 125

Punctation very dense and fine on frons and vertex, passing to
sparse on lower paraocular areas, sparser and surface more shining
than in barbota or jocasta, intervals between punctures equal to or
greater than puncture widths; clypeus and supraclypeal area a little
more shining with fewer punctures laterally; punctures of mesono-
tum and scutellum almost as dense as on frons, but on anterior
median part of former sparse, intervals between punctures dull
reticulate, over twice puncture widths; of mesepisternum above a
little coarser and slightly sparser than that of mesonotum, passing
to very sparse below; of metepisternum finer and shallower; sides
of propodeum a little roughly punctate, on postbasal angle and
posterior surface much less so; first two tergites entirely much more
finely punctate than mesonotum, distinctly sparser on first than on
second, third a little denser than second.

Head distinctly broader than clypeocellar distance (136:106);
upper interorbital distance longer than lower and equal to eye
length (84:78:84); interocellar distance clearly less than ocellocular
(20:25). Scape longer than half the maximum width of face (65:
49), surpassing the vertex; frontal carina interantennal, ending at
a point about half way from lower end to median ocellus; malar
space linear. Mesonotum anteriorly weakly produced medially,
pronotal corners forming obtuse angles with rounded apices; .
metanotum irregularly punctulate. Basal area of propodeum
broader than metanotum (26:19), not depressed in semilunar form,
but, with very short radiating basal carinae, and with reticulate
part forming cusp shaped figure.

Length 9 mm.; wing length including tegula 6.7 mm.; width of
head 2.3 mm.; of abdomen 3.1 mm.

Male: Color as in female, but with green spot at base of man-
dible, clypeus entirely green, scape with metallic reflections, flagel-
lum quite dark, especially first segment; posterior basitarsi dark
brown without metallic reflections.

Pilosity white, better developed than in female, but much less
than in males of other species.

Punctation as in female but a little denser on frons and
mesonotum, a little sparser on scutellum; on clypeus and supra-
clypeal area sparser than in female and intervals between punctures
shining; on episterna a little denser than in female; but punctation
little evident on sides of propodeum; on tergites a little stronger
than in female, but finer and sparser than on mesonotum, even on
sides of first tergite.

126 The University Science Bulletin

Head considerably broader than clypeocellar distance (140:110);
eye length greater than upper interorbital distance and this greater
than lower (90:85:68); interocellar distance almost equal to ocel-
locular (23:24). Scape slightly shorter than half of maximum
facial width (40:50) but clearly longer than first three flagellar
segments together, latter three segments of about same lengths
and each about as broad as long; malar area linear; frontal carina
vestigially indicated above interantennal tubercle; preoccipital
carina absent. Mesonotum with anterior part little produced, as
in female; femora and tibiae little swollen, even posterior pair.
Basal area of propodeum without semilunar depression, a little
longer than metanotum, carinae a little more evident than in fe-
male; metasoma with fourth sternite a little emarginate, fifth more
sclerotized medially and there depressed to form a canal that nar-
rows toward apical margin.

Length 9.1 mm.; wing length including tegula 7 mm.; width of
head 2.3 mm.; of abdomen 2.8 mm.

Distribution: Tandil, 250 meters altitude (type locality), No-
vember, 1954 ( F. H. Walz); other specimens, same data (Juan
Foerster). The locality is in the Province of Buenos Aires, Argen-
tina.

Types: Holotype, allotype, three female, and fourteen male para-
types in the Snow Entomological Museum, University of Kansas;
eight paratypes in the collection of the author in Curitiba, Brasil;
three paratypes in each of the following collections: United States
National Museum, American Museum of Natural History, British
Museum (Natural History), Museo Nacional "Bernardino Riva-
davia" (Buenos Aires), and that of Dr. Carlos Alberto Campos
Seabra (Rio de Janeiro).

Paroxystoglossa crossotos ( Vachal )

Halictus crossotos Vachal, 1904, Misc. Ent. [Narbonne], vol. 12, p. 118;

\'achal, 1911, Misc. Ent. [Narbonne], vol. 19, p. 14 (partini?).
Halictus (Conjnura'f) anthidioides Duckc, 1906 Zeitsclir. Hymenopterologic

Dipterologie, vol. 6, p. 397; Ducke, 1907, Zeitschr. Hymenopterologic

Dipterologie, vol. 7, p. 323.
Paroxystoglossa crossotos Moure, 1943, Rev. Ent. [Rio de Janeiro], xol. 14

p. 477 ( partim ) .

This species was described from "ouest de la Capitainerie des
Mines, Bresil" and the type is preserved in the Paris Museum. This
locality was presumably in Minas Gerais. Specimens studied by
Ducke were from Barbacena, in south central Minas Gerais. Vachal,
in 1911, probably included specimens of P. andromache under the

Review of the Genus Paroxystoglossa 127

name crossotos, as he referred to a specimen from Montevideo. I
have seen Ducke's type and have before me a specimen compared
with it.

Basically the two original diagnoses agree in principal features
that distinguish this species from its nearest relatives.

P. andromache is especially close to crossotos and was considered
a synonym by me in 1934. It now appears that they are different,
as indicated by the characters given in the key.

Distribution: Southern and eastern parts of Minas Gerais (Rar-
bacena) and eastern part of Sao Paulo (Serra de Bocaina) and
western part of Rio de Janeiro (Itatiaia).

Paroxystoglossa mimetica Moure
Paroxystoglossa mimetica Moure, 1950, Dusenia, vol. 1, p. 311.

The type locality is Onda Verde, Sao Paulo, Brasil.

The collection of the University of Kansas contains a specimen
from Rio Caraguata. Mato Grosso, Brasil, March, 1953 ( Fritz
Plaumann ) . This was compared with the type. It is slightly larger,
the wing (including tegula) measuring 6.2 mm. In this specimen
the greenish bronze reflection of the thorax and abdomen are even
less noticeable than in the type and the wings are a little more
strongly colored. The veins and hairs of the wing membrane along
the center of the wing are yellowish but veins G and R. the
pterostigma, the marginal cell, and hairs of the membrane near
these structures as well as on the wing open beyond the veins and
on the posterior border are dusky.

This species is easily separated from others of its group In the
lack of punctation on the first tergite, which is quite shining and
polished, with piliferous punctures very sparse on the basal third,
and with microscopic piliferous punctures on the apical depression
and on the lateral elevations.

Geographic distribution: Sao Paulo (Onda Verde) and Mato
Grosso (Rio Garaguata), Brasil.

Paroxystoglossa andromache ( Schrottky )

Oxtfstoglossa andromache Schrottky, 1909, Rev. Mus. La Plata, \()1. 16, p.
141; Schrottky, 1913, An. Soc. Ci. Argentina, vol. 75, p. 241.

Ualictus crossotos Vachal, 1911, Misc. Ent. [Narbonne], vol. 19, p. 14 (speci-
men from Montevideo only).

Paroxysioglossa andromache Moure, 1940, Arg. Zool. [Sao Paulo], vol. 2, p. 61.

Paroxijsioglossa crossotos Moure, 1943, Rev. Ent. [Rio de Janeiro], vol. 14,
p. 477 (part); Moure, 1950, Dusenia, vol. 1, p. 313.

The type locality is Puerto Bertoni, Paraguay. The allotype

was described from Curitiba and is preserved in my collection.

128 The University Science Bulletin

In the collections of the University of Kansas, Dr. C. A. Campos
Seabra ( Rio de Janeiro ) , the Departmento de Zoologia of the Sec-
retaria de Agricultura (Sfio Paulo), and the author are numerous
specimens of this species from Curitiba and Nova Teutonia. This
last locality is rather close to the type locahty.

It seems probable from the locality that Vachal's record of
crossotos from Montevideo relates to this species.

Some variations can be noted among specimens of this species.
The extent of the areas of yellow pilosity on the wing membrane
varies even among specimens from Nova Teutonia. In some the
yellow only includes the basal thirds of veins M + Cu and A and
adjacent wing membrane, while in others it extends to the separa-
tion of M + Cu from cu - a. In no specimen was any yellow ob-
served around the third submarginal cell.

Distribution: Parana (Curitiba, Ponta Grosso, Guarapuava),
Santa Catarina (Nova Teutonia) and Rio Grande do Sul (Esteio),
Brasil; Paraguay (Puerto Bertoni); and probably Uruguay (Monte-
video). It should also occur in Missiones, Argentina.

Paroxystoglossa seabrai sp. nov.

Female: Color black with faint metallic steel bluish reflections on
the clypeus, supraclypeal area, propleura, region of scuto-scutellar
suture, more greenish on fifth tergum and genal areas; on legs
reflections almost imperceptible. Wings fuscous, with vein and
hairs of area in middle of wing from base to a little beyond ends of
cells and between veins M and A (V) yellow; rest of wing and
veins dark, in contrast.

Pilosity largely white but shorter hairs fuscous, that on fore
and middle legs and hind basitarsi with fuscous hairs predominat-
ing; terga one and two without apical bands, that of third vestigial,
fourth a little more evident, these bands and all vestiture of fifth of
a slightly fuscous ferruginous; sternal pilosity long, white.

Punctation very dense and fine on frons, sparser and coarser
toward lower parocular areas, clypeus rather smooth with few
rather coarse punctures on the disc, denser and finer on upper
corners; finer and sparser on supraclypeal area, intervals on disc
dull, reticulate, three or four times as wide as puncture widths; on
mesoscutum very dense and fine, a little less on upper parts of
mesepisterna, passing to very sparse beneath, intervals there three
to four puncture wddths, dull reticulate, but smoother and wider
next to coxae; on metepisterna fine and dense; on sides of propo-

Review of the Genus Paroxystoglossa 129

deum with some granulate elevated punctures on a finely roughened
surface, posterior face of propodeum similar but more shining; on
first and second tergites fine and dense, a little finer and sparser
than on mesonotum.

Head distinctly broader than distance from clypeal margin to
anterior ocellus (125:95), upper interorbital distance shghtly
longer than inferior interorbital distance but shorter than eye
length (74:70:80); interocellar distance less than ocellocular (40:
55, ocellar diameter 25); scape longer than half width of face (60:
46), surpassing vertex; frontal carina clearly evident in lower three
fourths; anterior part of mesonotum projecting over pronotum, bi-
lobed, dorsolateral angles of pronotum salient, rounded, margin to
lobe gently concave; basal area of propodeum not defined, some-
what shining, reticulate, with short, very vague striae on basal
two fifths.

Size: Length 9 mm.; wing (including tegula) 6.7 mm.; head
width 2.2 mm.; abdominal width 2.5 mm.

Distribution: A^ude Solidao, Floresta da Tijuca, 500 meters, Rio
de Janeiro, D. F., Brasil (type locality), January, 1955 and 1956
( C. A. C. Seabra ) ; Vista Chinesa, Floresta da Tijuca, 600 meters,
February 23, 1956 (C. A. C. Seabra); Floresta da Tijuca, February
22, 1956 (C.A. C. Seabra).

Types: Holotype in the collection of Dr. Carlos Alberto Campos
Seabra, Rio de Janeiro; paratypes in the collections of the author,
the University of Kansas, the U. S. National Museum, and Dr.
Seabra.

Paroxystoglossa spiloptera sp. nov.

Female: Like P. seabrai but with the following differences:
Green reflections more accentuated on sides of face, upper part
of supraclypeal area, genal areas, pronotum, metanotum, mesepi-
sterna above; margins of tergites discolored, more broadly so
apically, with green reflections on posterior parts of third and
fourth terga and on entire fifth. Wings more hyaline than in
seabrai, veins and hairs fuscous except small area with yellow hairs
consisting of posterior internal two thirds of second and third
submarginal cells and adjacent region of second medial cell and
small area beyond second r - m, these together forming a distinct
contrasting spot; small area of yellow hairs at base of wing.

Pilosity denser covering corners and lobes of pronotum and meta-

5—8920

130 The University Science Bulletin

notum; tergal fasciae fuscoferruginous on terga two or four; pilosity
of fifth tergum fuscous.

Punctation as in seabrai but on clypeus and supraclypeal area
still sparser and their surfaces smooth and shining; mesepisterna
with intervals between punctures dull, reticulate; punctures of sides
of first tergite very sparse, almost absent, of disc very sparse with
intervals several times as great as punctures in contrast to dense
punctation of second.

Head distinctly broader than long (132:100); upper interorbital
distance slightly greater than lower which is equal to eye length
(86:78:78); interocellar distance distinctly less than ocellocular
(50:70, ocellar diameter 30); scape longer than half width of face
(60:50), surpassing vertex; frontal carina very short, only lower
third evident. Thorax as described for seabrai.

Length 7.5 mm.; wing (including tegula) 6.2 mm.; width of face
2.3 mm.; width of abdomen 2.5 mm.

Distribution: Guaruva, Santa Catarina, Brasil, 8 meters, October
30, 1955 (J. S. Moure, C. D. Michener).

Types: Holotype in the collection of the author, Curitiba, Brasil.
Ten paratypes distributed among the collections of the Snow En-
tomological Museum, University of Kansas; Carlos Alberto Cam-
pos Seabra, Rio de Janeiro; the U. S. National Museum, and the
author.

Specimens similar to those from Guaruva and probably belonging
to the same species have been collected in the Floresta da Tijuca
with specimens of P. seabrai.

Key to the Species of Paroxystoglossa

1. Males 2

Females 10

2. Second flagellar segment but little longer than first, distinctly shorter
than first and pedicel together; scape relatively long and slender,
almost reaching level of anterior ocellus; preoccipital carina absent 3
Second fiagellar segment distinctly longer than first, equal to or longer
than first and pedicel together; scape short and thick, terminating
more than its diameter below margin of anterior ocellus; preoccipital
carina distinct at least dorsally 4

3. Antennae relatively short, not reaching scuto-scutellar suture, flagellar
segments approximately as long as their diameters; horizontal part of
propodeum as long as or longer than metanotum with the depressed
area cusp shaped and with short basal carinae; punctation of frons
coarser than that of mesonotum, dense, but shining; upper interorbital
distance less than eye length; clypeus rather salient; head width httle

Review of the Genus Paroxystoglossa 131

greater than distance from clypeal margin to upper margin of anterior

ocellus ( 135: 120 ) brachycera

Antennae longer, surpassing scuto-scutellar suture, flagellar segments
longer than their diameters; horizontal part of propodeum very short,
depression semilunar, with radiating striae a little weak; punctation
of frons dense, dull, as coarse as mesonotum; upper interorbital dis-
tance longer than eye length; clypeus not very salient, but well ele-
vated toward apex; head width considerably greater than distance
from clypeal margin to upper margin of anterior ocellus (150:
120) harhata

Abdomen elongate elipsoid, wholly metallic, as are most parts of the
legs; femora and tibiae relatively swollen, at least posterior pair . 5
Abdomen decidedly claviform, largely dark, at least on tergal discs;
femora and tibiae normal 6

Tergites one to three with marginal depressions glabrous and smooth
in strong contrast to strong dense punctation of discs; punctation of
first tergum coarser than that of mesonotum, on terga four and five
considerably finer and very sparse; anterior and middle femora more
swollen than posterior; anterior tibiae metallic green, basitarsi dark
brovvTi transversa

Tergites one to three with piliferous punctures; punctation of first
tergite less strong than that of mesonotum, tergites four and five simi-
lar to preceding ones; posterior femora more swollen than anterior and
median ones; inner faces of anterior tibiae and all basitarsi pale yellow.

jocasta

Head above and entire thorax dark brown, without metallic colora-
tion or sometimes metanotum weekly greenish 7

Head and thorax entirely metallic green, sometimes a little dark on
discs of mesonotum and epistema 8

Median region of wing entirely covered with yellowish pilosity, veins
in this area also yellow; marginal depressions of terga broadly bleached
and covered with contrasting yellow hair [hypothetical characters for

the male which is unknown to me.] crossotos

Yellow portion of wing limited to basal third; abdominal bands, both
of bleached integument and of yellow pilosity, relatively narrow and
more or less vestigial on first two terga andromache

First tergite smooth and shining, with punctation very sparse (at the
sides punctures separated by over five times their diameters), on other
tergites punctation fine but dense; punctation of frons a little sparse,
a little coarser than that of mesonotum; epistema and sides of pro-
podeum reticulate but shining; size small, wing length (including
tegula ) 5.8 mm mimetica

First tergite rather densely punctate, intervals less than twice a punc-
tvire width; other terga densely and relatively strongly punctate;
epistema and sides of propodeum dull 9

132 The University Science Bulletin

9. Wing length (including tegula) 6 mm.; dorsum and sides of thorax
and upper part of head quite dark; anterior discal part of mesonotum
densely punctate, on scutellum rather sparsely punctate at sides of

median line spiloptera^

Wing length (including tegula) 7 mm.; dorsum and sides of thorax
and top of head clear green; anterior discal part of mesonotum less
punctate near midline; scutellum densely punctate seahrai^

10. Color predominantly metalhc, even on abdomen; abdomen subelipsoid, 11

Color dark on thorax and abdomen (see also some transversa), with
yellow pilosity more or less developed on margins of apical tergites;
abdomen subpedunculate; angles of pronotum more clearly marked,
as is anterior projection of mesonotum 14

11. Marginal depressions of tergites black, broadened to form black
bands especially on first two tergites, depressions glabrous, impunc-
tate, in contrast to strong dense punctation of discs of tergites where
it resembles that of mesonotum; basal area of propodeum not de-
pressed, and with fine transverse striae in the middle near posterior
declivity transversa

Marginal depressions of tergites similar to rest of discs, with sparse
piliferous punctures; punctation of discs of first two tergites con-
siderably finer than that of mesonotum 12

12. Vestiture of discs of tergites two to five consisting of pale or brown
hairs and short brown pilosity; scape metallic; tibiotarsal scopa vdth
pale or brown hairs jocasta

Vestiture of discs of tergites two to five entirely brown, as are hairs of
tibiotarsal scopa; scape nonmetallic 13

13. Mandibular base with metallic green spot; basal area of propodeum
shorter than metanotum, its posterior margin medially elevated as a
carina having the form of a very wide open V; punctures of first and
second terga rather dense, like those of third or denser barbatn

Mandibular base without metallic green spot; basal area of propodeum
longer than metanotum, grading into posterior surface without mar-
ginal carina; punctures of first tergite considerably sparser than those
of third hrachycera

14. First tergite almost entirely smooth, very shining, with only a few
fine, sparse, piliferous punctures in basal region and in marginal de-
pression mimetica

First tergite vidth at least apical third quite densely punctate, al-
though sometimes punctures fine and shallow and intervals between
them smooth and shining 15

15. Marginal depressions of tergites two to four broadly bleached and
covered with yellow appressed hairs, giving the appearance of bands
that occupy about one third of postgradular parts of terga; first tergum
generally vdth hair band narrower, although clearly visible 16

1. Males here characterized were collected with females, but as the two species occur
together there is no absolute certainty that they are correctly associated with the females
and I have not designated male allotypes.

Review of the Genus Paroxystoglossa 133

Marginal depressions of tergites one and two black, and generally with
only blackish hairs; third tergum narrowly pale, fourth more broadly
so and with yellow hairs forming a band 17

16. Forewings with veins almost entirely yellow and membrane with
yellowish hairs; veins C and R and pterostegina and anterior margin
of wing fuscous and with fuscous pilosity in marginal cell; apical mar-
gins of first four tergites with yellow pilosity well developed, especially

on two to four crossotos

Forewings generally with veins of apical half more or less dark, as is
the pilosity of the membrane in this area (sometimes only basal third
of wing, along the middle, with yellow); bands of terga less yellow,
that of first generally httle noticeable andromache

17. Wings vdth veins and hairs yellow along middle of wing between
M-Cu and A and including cells and veins demarking them, three
submarginal cells, the medias and cubitals, inner posterior side of
1st Ri and a little of tliird submarginal; rest of wing with veins and
hairs dark, in contrast seabrai

Wings with veins dark and hairs of membrane fuscus, except small
area with yellow hairs including inner posterior two thirds of second
and third submarginal cells and adjacent parts of second media and a
small part of 2nd r-m, forming a distinct spot contrasting vdth the
rest of the vdng spilaptera

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 6

Comparative External Morphology and Taxonomy of
Nymphs of the Trombiculidae (Acarina)^

D. A. Crossley, Jr.2

Abstract: Laboratory reared nymphs of 46 species of trombiculid mites
were used as the basis for morphological and taxonomic studies. The nymphs
are free-living and are rarely collected. Chigger mites, the larvae of trombic-
ulids, are commonly found as ectoparasites on many species of vertebrates.
Until recently the classification of these mites has been based almost entirely
on the larval stage.

Only a few of the known species were available as nymphs; these were
members of the genera Acomatucanis (2 species), Chatia (1 species), Hanne-
mania (3 species), Leeuwenhoekia (1 species), WJiartonw. (1 species),
Blankaartia (2 species), Cheladonta (2 species), Euschongastia (8 species),
Euschungastoides (2 species), Neoschongastia (2 species), Pseudoschbngastia
(2 species), Speleocola (1 species), Tromhicula (18 species), and Walchia
( 1 species ) . The morphological work was directed towards the discovery of
useful taxonomic characters. When discovered these characters were used to
provide diagnoses and keys for subfamilies, genera, subgenera, and species, in
a system essentially similar to those based on larvae. Few changes in the
classification were made; these were the suppression of the subfamily Walchiinae
as a synon>Tn of Trombiculinae, the elevation of Euschongastoides to generic
rank, and the transfer of Euschiingastia loomisi to that genus.

A comparison of the classification based on nymphs with that based on
larvae shows that most genera are distinct, i. e., recognizable by a character or
group of characters, in both larval and' nymphal stages. The degree of dis-
tinctiveness often is not the same in the two stages. No characters were found
in the nymphal stage which can be used to diagnose the subgenus Neotrom-
bicula (genus Tromhicula); however, relationships do not seem to be contrary
to those shown in the larval stage. In such cases the two systems of classifica-
tion are in basic agreement. A more serious discrepancy concerns the genus
Euschungastoides. Larvae of the two included species fall into two diflFerent
genera but the nymphs are similar to each other and are amply distinct from

1. Contribution No. 1031, Department of Entomology, University of Kansas.

2. Present address: Health Physics Division, Oak Ridge National Laboratory, Oak
Ridge, Tennessee.

(135)

136 The University Science Bulletin

other species. In this instance the difficulty appears to He in the use of a
single key character which separates two otherwise similar larvae into different
genera.

It is proposed that characters of postlarval stages should be considered as
additional evidence in the taxonomy of these mites, and that neither larval

nor postlarval stages should be given undue weight. While larva and post-
larva are adapted to different modes of existence there appears to be no base
for weighing characters of any particular stage.

TABLE OF CONTENTS

PAGE

Abstract 135

Introduction 137

Acknowledgments 139

Historical Account 139

Position of the Nymphal Stage in the Life History 140

Materials and Methods 141

External Morphology of the Trombiculid Nymph 143

Gross Morphology 143

Setae 144

The Gnathosoma 146

Basis Capituli 146

Hypostome 147

Chelicerae 148

Measurements of chelicerae 151

Palpi 153

The Scutum 162

The scutum in the Leeuwenhoekiinae 163

The scutum in the Trombiculinae 166

Measurements of the Scutum 172

The body 174

Shape 174

Size 175

Eyes 175

Setae 172

Sternum 178

Genitalia 179

Anus 179

Legs 180

Relations of Morphology to Habits and EN\aRONMENT 182

Comparison of Larval and Nymphal Morphology 184

Taxonomy of the Nymphal Stage 187

Family Trombiculidae 189

Subfamily Leeuwenhoekiinae 192

Genus Leeuwenhoekia 193

Genus Acomatacarus 195

Subgenus Acomatacarus 196

Subgenus Xenacarus 197

Genus Chatia 198

Genus Hannemania 200

Genus Whartonia 203

Nymphs of the Trombiculidae 137

PAGE

Subfamily Trombiculinae 205

Genus Blankaartia 208

Genus Cheladonta 211

Genus Euschongastia 215

Genus Euschongastoides 226

Genus Neoschbngastia 231

Genus Pseudoschbngastia 235

Genus Speleocola 238

Genus Trombicula 240

Subgenus Eutrombicula 244

Subgenus Neotrombicula 251

Subgenus Leptotrombidium 256

Subgenus Trombicula 258

Genus Walchia 268

Additional Genera Known as Postlarvae 271

Genus Ascoschongastia 271

Genus Doloisia 271

Genus Gahrliepia 272

Genus Guntherana 272

Genus Heaslipia 273

Genus Ipotrombicula 273

Genus Schongiistia 274

Genus Schoutedenichia 274

Genus Speotrombicula 274

Genus Walchiella 275

Comparison of Systems of Classification 275

Rudiments of Phylogeny 277

Literature Cited 281

INTRODUCTION

This paper presents the results of a comparative study of the
external morphology of the nymphal stage of the available species
of trombiculid mites, with the purpose of evaluating morphological
features as specific and as group characters, and presenting work-
able descriptions of the nymphs studied. This work is necessarily
of a preliminary nature and more detailed studies of some features
will no doubt produce further valuable information. An attempt
was made to include as many species as possible, but nymphs of
only a small fraction of the described species were available. More-
over, some of the species included are represented by but one or
two specimens. It is hoped that this study will provide a basis
for other workers in preparing descriptions and in discerning group
characters and features distinguishing the higher categories.

This study is restricted almost entirely to American species, for
practical reasons. Nymphs from other regions have been and are

138 The University Science Bulletin

being studied by other workers, including Audy in Malaya, Domrow
and Womersley in Australia, Lipovsky and others for Korea, Sasa
in Japan, and Vercammen-Grandjean in Africa.

The virtual restriction of this paper to one geographic area seems
to preclude any extensive revision of the extant classification based
on larvae, since the latter is based on the fauna of all regions. The
taxonomic effects of postlarval characters on the larval classifica-
tion is assessed but changes in the formal classification are kept to
a minimum.

The nymphal stage rather than the adult stage was selected for
study due to several considerations. The nymph difiFers from the
adult in only three major ways: it is of smaller size, has fewer setae,
and has incompletely developed genitalia (Wharton and Fuller,
1952:39). The reduced number of setae is actually an advantage
since study of variation of their number is greatly simplified. Pre-
liminary studies of adult genitalia failed to reveal characters of
value in distinguishing groups, although characters of this type
quite possibly are present. Thus little is lost in choosing the
nymphal stage rather than the adult. Moreover, nymphs may be
obtained from engorged larvae with relative ease, while the rearing
of adults requires not only suitable culture conditions but a food
material for the nymphs as well. The proper food for postlarval
stages is still a major deterrent in rearing trombiculids (Wharton
and Fuller, 1952:24) although much progress has been made
towards overcoming this difficulty (Lipovsky, 1954:945). By choos-
ing nymphs instead of adults, a greater number of species, repre-
sented by more specimens, was available for study.

In contrast to the larval stage, knowledge of the form and habits
of the free-living postlarval stages has only recently been acquired.
Although trombiculid adults are but rarely collected in nature, in
the past ten years rearing techniques have been devised so that
nymphs and adults may be reared from engorged larvae taken
from hosts. However, taxonomic descriptions of reared adults,
properly associated with their larval stage, have been slow to
appear in the literature, and most of the published descriptions are
somewhat superficial and inadequate for recognition of the species.
Accurate and complete accounts of postlarval stages are restricted
to a few species, but these have been made the basis of consider-
able generalization.

Nymphs of the Trombiculldae 139

ACKNOWLEDGMENTS

I wish to express my gratitude to those persons who contributed
ideas, efforts, and specimens to this work. I am particularly in-
debted to the staff and students of the department of Entomology,
University of Kansas, for such assistance.

To the members of the University of Kansas Chigger Project, I
extend my thanks for information, advice, and specimens. I am
especially grateful to Dr. Richard B. Loomis and Mr. Louis J.
Lipovsky without whose enthusiastic cooperation this work would
not have been possible.

For the direction of this work I thank Drs. Robert E. Beer and
Charles D. Michener, Department of Entomology, University of
Kansas. Also, I am grateful to Dr. Henry S. Fitch, Department of
Zoology, for critically reading the manuscript.

This work was supported by a grant (E-476R) from the National
Microbiological Institute, National Institutes of Health, U. S. Public
Health Service.

HISTORICAL ACCOUNT

Th history of the development of knowledge of trombiculids has
ben reviewed in several recent publications (Wharton and Fuller,
1952; Womersley, 1952, and others); an account of recent develop-
ments should suffice for the purposes of this paper.

Three major evaluations of the Trombiculidae have been pub-
lished recently, all of which consider both larvae and postlarvae.
Wharton and Fuller (1952) presented a classification based on
larvae; but it utilizes some postlarval features and includes some
partial postlarval diagnoses. This effort summarizes our knowledge
of trombiculids and includes detailed synonymies for each species.
Womersley (1952) revised the Trombiculidae (of Womersley,
Trombiculinae of others ) by the use of postlarval evidence. Wom-
ersley places greater emphasis on postlarval stages than on larvae
and deduces from postlarval evidence that some larval characters
which were formerly given generic value may be of no more than
specific significance. Audy ( 1954 ) discusses characters of both lar-
vae and postlarvae, and comments on various genera. These com-
ments are based largely on postlarvae studied by him. Unfortu-
nately the descriptions of the nymphs mentioned have not yet been
published. Audy steers a course between the first two authors cited
and apparently has adopted the better features of each.

These papers indicate a trend towards a biologically realistic
classification of trombiculids, through a consideration of the char-
acters provided by all stages of the life history.

140 The University Science Bulletin

POSITION OF THE NYMPHAL STAGE IN THE LIFE

HISTORY

The stages in the life history of trombiculid mites are egg, deu-
tovum, larva, prenymph (nymphochrysalis), nymph, preadult
(imagochrysalis), and adult. The larva, nymph, and adult are
active stages; the egg, deutovum, prenymph, and preadult are
inactive stages in which development occurs. The larvae are
parasitic upon terrestrial vertebrates. The nymphs and adults are
free-living and predaceous upon small arthropods and their eggs.

The nymph is the first free-Hving active stage in the life history.
If it cannot endure the physical conditions of the habitat in which
it finds itself, or cannot find or compete for food materials, it must
eventually perish. The adult, also free-living, probably has much
the same requirements as the nymph (laboratory cultures so indi-
cate). Characters of the free-living stages may be considered to
be tested first by selection in the nymphal stage and those indi-
viduals insufficiently adapted to the particular environment never
reach the adult stage and hence never reproduce.

It has been shown that trombiculid larvae, while not host-specific,
may occur in greater abundance on one or a few particular hosts.
For example, Loomis (1955:255) shows that Trombicula giimeyi
larvae are found on several reptilian and mammalian hosts, but
are found with greater frequency and in greater numbers on those
closely associated with decaying logs. The adults have been found
in decaying logs, and unattached larvae have been taken from this
habitat. The skink, Etimeces jasciatus, is closely restricted to this
habitat, and consequently a greater percentage of larvae, which
detach from this host will find a more suitable habitat, than will
those which detach from the rattlesnake, Crotalus horridus, not so
associated with decaying logs. However, the larvae can apparently
engorge themselves successfully on either of these two hosts or on
others.

Thus the larval stage seems to serve primarily as a means of dis-
persal. The nymphal and adult stages might be considered as an
"anchor" which binds the species to a particular ecological situa-
tion and the larva to a particular set of host relationships.

From these considerations it appears that the nymphal stage is
at least as critical as the other stages of the life history. Taxonomic
characters of this stage should be as valuable and as stable as the
characters of the larva and adult. Of course, a sound classification

Nymphs of the Trombicxjlidae 141

must admit taxonomic characters of all stages. It is not proposed
that nymphal features should be given more weight than those
of any other stage,

MATERIALS AND METHODS

The specimens used in this work were mostly reared from en-
gorged larvae taken from vertebrate hosts. A few n^nnphs were
collected from soil, rodent nests, decaying wood, or from beneath
rocks. Methods used in the collection of host animals and in re-
moval of chiggers have been described in detail by Loomis
(1956:1218).

Living engorged chiggers were placed in culture dishes or tubes
lined with a mixture of charcoal and plaster of Paris (Lipovsky,
1953:4). Nymphs were preserved several days after emergence,
and were identified from larval specimens associated with them.
In later stages of the work, isolation tubes were found to be valu-
able. These were made from 1-dram vials by addition of a small
amount of the charcoal-plaster of Paris mixture. Inactive engorged
larvae were transferred singly from larger culture vials into these
isolation tubes. After the nymph had emerged, the larval skin
could then be mounted for identification of the specimen. The
skin was usually found without difficulty.

Specimens were mounted on standard microscope slides for
study. Polyvinyl alcohol with lacto-phenol was used as a mounting
medium and was found to be satisfactory; however, preliminary
clearing of the specimen in hot lacto-phenol produced better slides
than did mounting directly from alcohol. Mounts of living ma-
terial were usually inferior to mounts of material preserved in
alcohol for several days.

A phase-contrast microscope was used for detailed study of the
specimens. Drawings were made with the aid of a camera lucida.
Measurements of structures were made with an ocular micrometer
of the fixed type.

In the preparation of mites for taxonomic study customary pro-
cedure includes clearing of the soft internal parts so that only
skeletal structures remain. This procedure was followed in this
study so no description of the soft anatomy can be given. The
account is not entirely restricted to external anatomy, however,
since the few sclerotized internal structures remain after clearing.
Such internal structures, associated with the gnathosoma and geni-
talia, are included in the following discussions.

142 The University Science Bulletin

Nymphs of forty-six species of trombiculids are included in this
paper. A list of these species is presented below, modified by some
changes in classification. In the ensuing discussions the evidence
for these changes will be presented. It is somewhat less than logical
to present the classification and the changes before presenting the
evidence for them, but this course may be justified by the saving
of space and by the continuity obtained.

Unfortunately some species are represented by only one or a
few specimens. It is hoped that the knowledge of variability
gained from longer series of other species has enabled the author
to avoid gross errors in characterizing these forms.

No specimens of members of the subfamily Apoloniinae were
available for study, so that this group is necessarily excluded from
the discussions.

Family Trombiculidae Ewing

Subfamily Leeuwenhoekiinae Womersley
Acomatocartis (Acomatacanis) orizonensis Ewing
Acomatacorus (Xenacarus) phimostts Greenberg
Chotia setosa Brennan
Hannemania dtinni Sambon
Hannemania eltoni Sambon
Hannemania multifemorala Loomis
Leeuwenhoekia (Comatocariis) americana (Ewing)
Whartonia senase (Greenberg)

Subfamily Trombiculinae Ewing
Blankoartia alleei (Ewing)
Blanhaartia velascoi (Boshell and Kerr)
Cheladonta micheneri Lipovsky, Crossley, and Loomis
Cheladonta ouachitensis Lipovsky, Crossley, and Loomis
Euschongastia criceticola Brennan
Euschongastia cynomijicola Crossley and Lipovsky
Euschongastia diversa Loomis
Euschongastia jonesi Loomis
Euschongastia peromysci (Ewing)
Euschongastia pipistrelli Brennan
Euschongastia setosa (Ewing)
Euschongastia trigenuala Loomis
Euschongastoides hoplai (Loomis)
Euschongastoides loomisi (Crossley and Lipovsky)
Neoschongastia americana (Hirst)

Nymphs of the Trombiculidae 143

Neoschongastia brennaiii Crossley and Loomis

Psetidoschongostia farneri Lipovsky

Pseudoschongastia htmgerfordi Lipovsky

Speleocola tadaridae Lipovsky

Trombictda (Etdwmhictda) alfreddngcsi (Oudemans)

Trombictda (Etitrombicula) belkini Gould

Trombictda (Entrombictda) lipovskyona Wolfenbarger

Trombicida (Etitrombicula) splcndens Ewing

Trombictda (Lepfotrombidiiim) mtjotis Ewing

Trombictda (Neotro7nbictda) atitwnnalis (Shaw)

Trombictda (Neotrombictda) Upovskyi Brennan and Wharton

Trombictda (Neotrombictda) sijJvilagi Brennan and Wharton

Trombicida (Neotrombictda) tchartoni Ewing

Trombictda (Trombictda) crossleyi Loomis

Trombictda (Trombictda) fitchi Loomis

Trombictda (Trombictda) gurneyi Ewing

Trombictda (Trombictda) kansasensis Loomis

Trombictda (Trombictda) kardosi Loomis

Tro7nbictda (Trombictda) merriheivi Loomis and Lipovsky

Trombictda (Trombictda) montanensis Brennan

Trombictda (Trombictda) ornata Loomis and Lipovsky

Trombicida (Trombictila) trisetica Loomis and Crossley

Walchia americana Ewing

EXTERNAL MORPHOLOGY OF THE TROMBICULID

NYMPH

Gross Morphology

The generalized form and structure of trombiculid nymphs have
been known since the pioneer work of the Japanese investigators.
Hirst (1925:609) gives an excellent figure of the audit of Trombic-
ula autumnalis and Andre (1930) gives a clear account of the
morphology of that species. Other illustrations and descriptions of
some postlarvae were published by various workers prior to and
during World War II. Some of these contain inaccuracies since
the figures were intended to present the generalized form of the
animal rather than exact detail. In 1946 Wharton described the
sclerotized anatomy of all stages of Etischongastia indica, with
attention to detail. Michener (1946a), Jenkins (1949a), Richards
(1950), Wharton et al (1951), Wolfenbarger (1952), Wharton
and Fuller (1952), Womersley (1952), Brown (1952), Sasa (1953)
and others have described morphological features of postlarvae.

144 The University Science Bulletin

The trombiculid nymph is an eight-legged form with a pair of
chelicerae and a pair of palpi. Primary segmentation is obscured.
The body may be divided into four general regions, tlie gnatho-
soma, the propodosoma, the metapodosoma, and the opisthosoma.
The gnathosoma is the region bearing the mouth and mouth parts
while the propodosoma bears the first two pairs of legs, the meta-
podosoma bears the second tu'O pairs of legs, and the opisthosoma
is the remaining posterior portion of the abdomen. In most trom-
biculids the propodosoma and metapodosoma are separated dor-
sally by a strong constriction, the body anterior to the constriction
being called the proterosoma and that posterior to the constriction
the hysterosoma. The gnathosoma is the most distinct region, and
sometimes the entire body posterior to the gnathosoma is called
the idiosoma.

The idiosoma has a single dorsal plate, the scutum or crista
metopica, on its anterior dorsal portion. Ventrally, the area be-
tween the coxae of the first two pairs of legs is called the sternum.
The genital opening and the anus are found on the venter between
or behind the coxae of the last pair of legs. One or two pairs of
eyes, when present, may be found on the anterior dorsal aspect
of the idiosoma.

The four pairs of legs are borne in two groups, two pairs ante-
riorly and two posteriorly. The tarsi of all legs are provided with
claws. All of the legs are used in walking, although the first pair
of legs evidently functions as a sensory organ as well.

Setae

Numerous and highly modified body setae characterize the
group of prostigmatic mites to which trombiculids belong. Setae
are also numerous and somewhat modified on the appendages. In
the family Trombidiidae the forms of the body setae have been much
used as generic and specific characters; but in the Trombiculidae,
while these setae are not so strongly modified they are still useful
as taxonomic characters. The specialized setae of the body and the
scutum will be discussed in detail in other sections of this paper
but the appendicular setae merit some special consideration.

In the classification of trombiculids based on larvae, the form
and position of setae have been used as specific and as group char-
acters. Wharton et al. (1951:13) standardized a system of termi-
nology for the appendicular setae which has been widely accepted.
The appendicular setae of larvae and postlarvae are essentially
similar, so that some features of the terminology for larval setae

Nymphs of the Trombiculidae 143

may be adapted for postlarvae. This scheme has not been fol-
lowed invariably, since it seems preferable to propose a new term
rather than to misapply the larval terminology.

The setae found on trombiculids fall readily into two classes:
thick- walled branched and unbranched (nude) setae, and thin-
walled nude setae. Thick-walled setae usually consist of a long
stem with an attenuated tip and have few to many shorter branches
which arise at more or less regular intervals, and often from sev-
eral planes. Thin-walled setae are typically shorter, rather thick-
ened, pointed or blunt at the tip, and are often transversely stri-
ated. Specializations of both types occur but the only point of
confusion concerns thick-walled or "branched" setae which have no
branches. Jones (1950:486) described in detail sectioned setae of
the larva of Trombicula autumnalis and demonstrated that branched
setae (both with and without branches) have thick walls while
nude setae ("peg organs") have thin walls. Grandjean has called
the thin-walled setae "solenidions." The thin-walled condition is
evident under the phase-contrast microscope without special prep-
aration, since an internal cavity can be seen in the basal portion of
these setae; such a cavity is visible only in some of the larger
branched setae and here the thick-walled condition is obvious.
Thus in practice the separation of the two types usually presents
no difficulties.

In postlarvae, thin-walled nude setae occur only on the append-
ages, these setae being increasingly numerous on the leg segments
distal to the tiochanter, and one such seta occurs on the palpal
tarsus. All of the setae found on the body, including the nude setae
on the hypostome, those on the genitalia, and the sensilla of the
scutum, appear to be thick-walled setae derived from the branched
type. The larval system for naming the thin-walled setae involves
combining the name of the segment on which the seta occurs with
the suffix "-ala." Thus, a thin-walled nude seta found on the tibia
is called a "tibiala." For postlarvae, the terms basifemorala, telo-
femorala, genuala, tibiala, and tarsala may be used for these setae.
Such terminology is used throughout this paper, the term "nude
seta" referring only to thick-walled, unstriated setae that lack
branches.

Many of the appendicular setae are set in pits, so that it is usu-
ally difficult to decide whether a circular structure surrounding the
base of a seta is a pit or an alveolus. In the illustrations an alveolus
is shown for each branched seta but is omitted from the thin-walled
nude setae. This is probably the condition which obtains.

146 The University Science Bulletin

The Gnathosoma

The gnathosoma is the most anterior and the smallest division
of the body. It is completely fused to the propodosoma, with lit-
tle trace of division ventrally and none dorsally. The gnathosoma
consists of a somewhat conical basal portion (basis capituli) bear-
ing the palpi dorsolaterally and the chelicerae dorsally, and a pro-
jecting distal portion (hypostome). These parts are discussed sep-
arately below.

The morphology of the gnathosoma of the adult of Trombicula
alfreddugesl has been well described by Brown (1952), and his
descriptions appear to be generally applicable to both nymphs and
adults of most species. Brown's work is a morphologic and not a
taxonomic effort. His descriptions are difficult to apply in some
instances to specimens mounted on slides in the usual manner.
Thus discrepancies between Brown's presentation and that of the
writer are partly due to differences in purpose and method of study.

Basis capituli. The exact extent of the basis capituli and its
connection to the propodosoma are obscured by the process of
clearing and mounting. The sclerotized structures mentioned by
Brown (1952) may usually be located although no longer in
normal position. Most of these sclerotized structures form a cap-
sule (essentially the basis capituli) which is but rarely broken in
mounting and is occasionally separated completely from the body.
Apparently this capsule consists of the structures which Brown has
termed epistome (E), epistomal apodemes (EAs), and apodeme at
the junction of the basis capituli with the propodosoma (BCA).
The hypostome is found at the distal end of this capsule, and oc-
casionally the palpi remain completely articulated after mounting.
The chelicerae are almost invariably displaced and the integumen-
tary connections of the capsule with the body are either destroyed
or obscured in mounting. The capsule (fig. 198) is roughly coni-
cal in shape, open at both ends and open dorsally, the two sides
being connected dorsally by a single sclerotized crosspiece near
the anterior end. The posterior margin, the dorsal margins, and
the dorsal portions of the anterior margin are sclerotized. Appar-
ently the dorsal portion behind the crosspiece is internal. The
depressed space anterior to the crosspiece contains the chelicerae
and the palpi are articulated dorsolaterally on the anterior margin.
The hypostome projects from the anterior ventral portion.

The shape of the capsule in the species studied is approximately
the same as that of Trombicula trisetica (fig. 198). The capsule is

Nymphs of the Trombiculidae 147

somewhat broader and appears shortened in Acomatacarus plu-
mosus, Euschongastia jonesi, E. pipistrelli, species of Euschongas-
toides, Hannemania eltoni, and species of Pseudoschongastia; it is
narrowed and somewhat elongated in Speleocola tadaridae.

The chehceral apodemes ( CA ) and the skeletal structure forming
lateral articulations for the chelicerae (ChA) appear to be firmly
connected and usually remain so after mounting although no longer
in position.

The pharynx is a broad, flattened tube which can be seen occa-
sionally in the gnathosoma of mounted specimens. However it
was not seen in all of the species examined. In certain species
(Eiischdngastoides, Psetidoschongastia, and Walchia americana)
the pharynx is quite distinct and is marked with a reticulate pattern
(fig. 8). The significance of this marking is not known. In Pseudo-
schongastia hungerfordi the pattern is restricted to the midline of
the pharynx; in the other species the reticulations cover about half
the width of the pharynx.

Hypostomc. In mounted material the hypostome appears as a
lobe projecting from the ventral part of the basis capituli; in un-
mounted specimens it appears coneshaped with the dorsum open.
Mounting distorts the shape of the hypostome but it usually ap-
pears triangular and continuous with the venter of the basis ca-
pituli. The ventral portion bears setae; the dorsal portion bears a
sclcrotized hypostomal trough (Brown, 1952).

In most species the hypostome appears triangular and bears about
eight nude setae, usually four on each side at the apex (fig. 9).
Posterior to the nude setae are about ten to twenty branched setae,
similar to those on the venter of the basis capituli. Usually it is
impossible to delineate clearly the venter of the hypostome from
that of the basis capituli, so that some setae cannot be assigned
accurately to either. There is some intraspecific variation in the
number of nude setae, occasional specimens having as many as ten
or as few as six. These setae occasionally bear one or two fine
branches. There are some marked and consistent variations from
this typical condition; these merit further characterization.

In Eiischdngastoides, Pseudoschongastia, and Walchia americana
the hypostome is blunt, and occasionally the margin is slightly
concave (fig. 8). Along the margin are about twenty short setae
which are nude or have one or two branches. Behind these, some
twelve longer branched setae are distributed over the venter of the
hypostome. In Walchia americana the edge of the hypostome is

148 The University Science Bulletin

not as straight as in the other species; usually it is slightly convex.

Species of Cheladonta have the hypostome more elongated than
do the typical species of the family and the apex bears eight to ten
rather short nude setae (fig. 10). About ten nude setae are dis-
tributed over the remainder of the hypostome; occasional setae have
one or two branches but there are no heavily branched setae on
the hypostome.

Species of Neoschongastia have a rather blunt hypostome which
bears eight short nude setae on its apex (fig. 11); some ten branched
setae are scattered over the more posterior portion. The apical
nude setae are very short in N. brennani and somewhat longer in
N. americana.

Acomatacarus phimoms and Wliartonia senase have a blunt
hypostome which bears six to eight apical setae, all of which have
several branches. About twenty slightly expanded, branched setae
are found on the remainder of the hypostome.

In Chatia setosa and species of Hannemania the branched setae
covering most of the hypostome are quite numerous. Some of the
specimens of Hannemania eJtoni have as many as fifty such setae.

Chelicerae. The chelicerae are composed of two segments; a
proximal elongated base and a distal blade. The base is articulated
to the basis capituli and has a slight amount of forward-backward
movement. The blade is articulated with the base at the sides
to allow for dorsal-ventral movement. The chelicerae have no
setae.

In mounting, chelicerae usually become partially detached and
are seen in side view. This position seems to be favorable for
characterizing the base and it is certainly the best position for
observation of the blade. All of the drawings of the chelicerae
are made from the medial (inner) surface, and all measurements
are made in side view.

The base is typically elongated, roughly oval in cross section,
pointed posteriorly, blunt anteriorly, with a straight ventral margin
and an arched dorsal margin. The area of articulation with the
gnathosoma is on the posterior ventral portion and occupies about
one half of the length of the ventral portion, extending from the
posterior tip forward (see figures). In mounted material the area
of articulation is seen to be more on the medial surface than di-
rectly ventral (see also Brown, 1952, p. 47, fig. 4; p. 49, fig. 8).
The base is unusually elongated in Blankaartia, Cheladonta, Speleo-
cola tadaridae, Trombicula ornata, and T. kansasensis; the area of

Nymphs of the Trombiculidae 149

articulation is also elongated in S. tadaridae, T. kansasensis, and T.
ornata. The articulating area appears to extend across the posterior
part of the base instead of the ventral surface in Eiischongos-
toides, Pseudoschongastia, and Walchia americana (figs. 30-33, 35).
The area of articulation appears shortened in Neoschongastia and
Whartonia senase. Otherwise, characterization of the shape of
the base is difficult, since individual variation, although slight, is
noticeable. In some species the base appears to have a more robust
shape, i. e., shorter and broader, than in others. However, diflFerent
specimens of the same species present slightly diflFerent aspects
and intermediate conditions defy grouping. Aside from the obvious
deviations given above, the shapes of the cheliceral bases are more
or less similar.

Punctation is generally present on the cheliceral base, and ap-
pears to be more dense on the dorsal and perhaps the ventral
portions than on the lateral portions. Puncta are difficult to see
in poorly mounted specimens and even under good conditions
estimates of size and number of the puncta are difficult. Occasional
specimens and some species of specimens appear to lack puncta
altogether, and specimens preserved soon after they enter the
nymphal stage may lack them. Species which have punctation of
the cheliceral bases sparse or absent include Cheladonta ouachiten-
sis, Euschongasfia criceticola, E. cijnomijicola, E. pipistrelU, Eu-
schongastoides loomisi, Pseudoschongastia fanieri, P. hungerfordi,
Trombictda crossleyi, T. gtirncyi, and T. ornata. Puncta are par-
ticularly numerous on the cheliceral bases of Euschongastia diversa,
E. jonesi, E. peromijsci, E. trigenttala, and Hannemania dunni.
Conspicuously coarse punctation is present on the bases of Hanne-
mania dunni, Leeinvenhoekia americana, Trombictda sijlvilagi, and
T. whartoni.

The cheliceral blade usually b^ars a resemblance to a single
blade of a pair of shears, so the term "shcarlike" has been used in
the section on taxonomy to designate this usual condition. The
blade is flattened laterally, with an attenuate tip ending in a sharp
point and a row of teeth on the dorsal edge. The blade appears
to be adapted for piercing rather than for cutting. The proximal
portion of the blade is an upright structure, the apodeme of the
blade, lying partly within the cheliceral base. The blade articulates
with the base at two points, one on each side of the apodeme of
the blade. Apparently muscles attach here also. This apodeme is
slightly constricted in most species. The constriction is particu-

150 The University Science Bulletin

larly noticeable in species of Cheladonto, Euschongastia trigenuala,
Tromhiciila kansasensis, and T. ornata. No constriction was seen
in Acoinatacarus, Chatia setosa, Euschongastoides, Hannemania,
Tromhic.ula myotis, and Whartonia scnasc.

The distal portion of the blade in most species curves upward to
an acuminate apex. The blades of some species appear to be
broader than others; but as with the shape of the cheliceral base,
this character proved very difficult to use. Particularly broad, i. e.,
deeper and coming to a more abrupt point, are the blades of
Euschongastia criceticola, E. diversa, E. Peromysci, E. trigenuala,
Trombictda lipovshji, and T. sylvilagi. Related species approach
this condition to such an extent that the characterization is some-
what arbitrary (see illustrations). The few following types are
sufficiently distinct from the usual blade type to merit description:
Whartonia senase and Neoschongastia spp. have a considerably nar-
rowed blade which curves up at the tip; the blade of Acomatacarus
arizonensis is also narrowed but is straight at the tip, being dagger-
like; in species of Cheladonfa, the blade is broad but also strongly
curved. Elongations of the usual shape also occur; but these are
difficult to single out. However, species of Blankaartia seem to
show the greatest elongation,

A dorsal row of teeth on the cheliceral blade was found in all
species except Neoschongastia hrennani, which lacks teeth alto-
gether. It is possible that the two specimens of N. hrennani are
anomalous in this respect. Occasional specimens of other species
were found to lack teeth on one of the blades or to have the num-
ber greatly reduced. Often, but not always, it appeared that
these specimens had been preserved soon after emergence.

Cheliceral teeth vary in size and number among species, within
the species, and on the individual specimen. Customarily the
largest teeth are found near the base of the blade and become pro-
gressively smaller towards the tip, where they are tiny. This is
not a regular gradation, however; very tiny teeth frequently occur
at the base of the blade, and the row of teeth is usually uneven,
with teeth of various sizes occurring together. Some of the config-
urations suggest that a large tooth may break ofi^, leaving several
small points. The irregularity, together with the small size, makes
cheliceral teeth difficult to count and somewhat difficult to charac-
terize for a given species. In most instances there is a relationship
between the size and the number of teeth, species with larger teeth
having the smaller number, if such factors as the size of the mite

Nymphs of the Trombiculidae 151

or peculiar modifications do not disturb the comparison. This re-
lationship is less noticeable in variation within the species; individ-
uals with different numbers usually have teeth of similar size.
Counts of teeth were attempted for all species and these figures
are given in the descriptions as approximations. The character of
many small versus few large teeth has some value in separating
some of the species, but cannot be rigidly applied due to the in-
accuracies resulting from difficulty in counting. As noted above,
occasional specimens have a blade with the teeth greatly reduced
or absent.

Teeth are particularly numerous on the chelicerae of Chatia
setosa, Euschongastia pipistrelli, and E. sctoso. Large teeth, re-
duced in number, are found in Cheladonta micheneri, Pseudoschon-
gastia fameri, and Trombiciila sylvilagi. These two groupings in-
clude extreme cases; other species show tendencies toward these
extremes. Species of Cheladonto, Euschdrm.astoides loomisi, Whar-
tonia scnase, and to some extent E. hoplai, show a restriction of the
teeth to the distal portion of the blade. The row of teeth (the
dorsal margin of the blade) is straight rather than curved upwards
in Acomatacanis phimosus, Euschongastoides, Hannemonia multi-
femorala, and Whartonia senase; however, other species approach
this condition.

A membranous structure, the pseudochela, is usually visible as
a projection from the dorsal margin of the end of the cheliceral base.
This structure has been omitted from some of the illustrations but
it is believed to be present in all species although sometimes
difficult to see. Possibly it is a remnant of the fixed chela, or simply
an extension of the articulating membrane of the base-blade union.
The shape is variable, probably being influenced by mounting tech-
niques.

The cheliceral apodemes are visible in most specimens and are
generally similar in shape. Specific differences may exist in the
shape of the structure but if so they are masked by individual varia-
tion. These structu-es are also frequently distorted during the
mounting process. They have not been figured for most species.

Meastirements of chelicerae. Since the chelicerae usually fall
into side view during the mounting process, and are rarely distorted
by mounting, they were considered to be suitable subjects for
measurement. It was hoped that a measurement could be found
which would serve as an index of total body size, since this is not
suitable for accurate measurement due to distortion of the bodv in

152 The University Science Bulletin

mounting. Three measurements were made on the chehcerae, as
follows: BL: Length of the cheliceral base, measured from the
posterior extremity to the point of articulation of the blade on the
medial surface. BH: Height of the cheliceral base, measured at
the widest point. CL: Length of the cheliceral blade, measured
from the tip to the back of the apodeme of the blade.

Most of the samples of nymphs are small. Large samples (more
than thirty individuals) with identical data were available for
Trombicula lipovskyi and T. montanensis. Graphic checks for nor-
mality showed that the three measurements were skewed to the
right in T. lipovskyi (an abundance of larger values and a shortage
of smaller ones ) ; in T. montanensis they were normally distributed
or nearly so. Two ratios were made from the values, these being
BL/BH and BL/CL; these ratios were checked by graphic meth-
ods and found to be normally distributed in both species.

It should be emphasized that these measurements were taken in
an endeavor to discover taxonomic characters. It is to be expected
that two species could differ in a measurement but it is not par-
ticularly surprising when they do not; also, the figures given are
based on individuals which in some cases are probably members of
different populations of the species. This work is intended to show
that measurements of the chelicerae are practical for taxonomic
purposes, including investigations of variation within a species;
that these measurements in some cases may serve as characters for
the identification of species; and that the ratios used have a value
similar to that of the measurements on which they are based. The
measurements are used, however, only to separate species or groups
of species from one another where such separation is clear. The
ratios serve particularly to quantify relationships which are already
obvious.

Means of the three measurements and their ratios, with their
standard errors, are given in the species descriptions in the taxo-
nomic portion of this paper. Examination of these will show that
some of the species may be separated by cheliceral measurements
even though some accuracy is lost because of skewness. Also, the
measurements seem to give an indication of the general size of the
mite. Within a species group where cheliceral shapes are similar,
these measurements more accurately indicate relative size.

The ratios based on these measurements are not valuable in
separating closely related species from one another. However,
several interesting groupings of species appear, as shown in Tables
1 and 2. These ratios give a quantitative expression to some pe-

Nymphs of the Trombiculidae 153

culiarities evident in the illustrations, as far as the extreme forms
are concerned. That related species have similar ratios may be
seen in the Tables.

The ratio BL/BH ranges from about 1.8 to about 4.0. Most of
the species have a mean ratio between 2.3 and 3.0; the larger mean
ratios of Speleocola tadaridae (3.5), Trombictda ornata (3.6), and
species of Chchdonta (4.0) are distinctive. There is also a group
of species with mean ratios between 1.8 and 2.1; this group is not
so distinctly separated from the majority but is worthy of mention.
The group includes Acomatacarus arizonensis, A. phimosiis, and
Whartonia senase (1.8); Leetiwenhoekia americana and Psetido-
schongastia farneri (1.9); P. hiingerfordi, Euschongasfoides hophi,
and Walcliia americana (2.0); and E. loomisi and Hannemania
multifemorala (2.1).

The ratio BL/CL ranges from about 1.6 to about 3.7. The only
species clearly separated from the rest are members of the genus
Cheladonta (3.6 and 3.7).

Palpi. The palpi are the second pair of appendages of the
gnathosoma; they are attached anteriorly at the sides of the basis
capituli and project forward in front of the hypostome. The palpi
are arched dorsally, so that the anterior ends point downwards.
The segments are hinged on their dorsal margins. Brown (1952:
43) states that none of the muscles of the palpi are extensors. Five
free segments are recognizable in the palpus, these being trochanter,
femur, genu, tibia, and tarsus. The coxae are incorporated into
the basis capituli and are not apparent as segments. The palpus
has a chelate apex ("thumb-claw process" of authors); this con-
sists of a "thumb," the palpal tarsus, which articulates near the base
of the tibia instead of at the apex, and a claw arising from the apex
of the tibia. The chelate apex of the palpus is characteristic of
several families of prostigmatic mites.

The most favorable position for observation of the palpi is a
side view, so that the medial (inner) and lateral (outer) surfaces
are prominently displayed, and the dorsal and ventral surfaces
form the margins. The palpi usually fall into this position in
mounting.

In general, palpi of the different species have relatively similar
aspects. Some appear more elongated than others but a gradation
occurs. The short, broadened palpi of NeoscJiongastia appear to
be distinctive in shape but others (Pseudoschongastia) approach this
condition. General palpal shape within a species is sufficiently

154

The University Science Bulletin

Table 1. The Ratio BL/BH (Cheliceral measurements):
Species among Taxonomic Categories.

Distribution of

Mean Ratio

«
o

c
o

o
is

-*•

S
0

•S 1

1.8

3
1

1.9

2.0

2.1

2.2

2.3

2
1

"2"

1
1
1
1

2.4

"l
1

1
2

2.5

2.6

2.7

2.8

2.9

3.0

3.5

1 (Speleocola)
3)

3.6

1 (7". ornat

4.0

2 (Cheladonta
spp.)

Table 2. — The Ratio BL/CL ( Cheliceral measurements ) : Distribution of
Species among Taxonomic Categories.

Mean Ratio

S
0

&->

a>
0

.si

1.6

1
2

"2"
I

1
1
2

1.7

1.8

2
3

1
2

1.9

2.0

2.1

2.2

1
2

2.3

2.4

2.5

2.6

2.7

2.8

1 (T- nrnn.t'n.')

3.0

3.6

1 (Cheladonta

micheneri)
1 (C. ouachitensis)

3.7

Nymphs of the Trombiculidae 155

constant, however, that some species may be classified as having
palpi "elongated" or "shortened." These terms are subjective and
a more valuable characterization of the palpus requires a more
detailed discussion.

Certain features of the palpal setae require special consideration.
Except for specialized setae on the tarsus, palpal setae may be of
two types: unipectinate and bi- or multipeclinate. Unipectinate
setae are straight setae with branches arising from one side only.
The branches are thick basally, taper abruptly, and run parallel
to the body of the seta. Other setae are flexible; the branches
taper gradually, run in different directions, and arise from more
than one side of the seta. These flexible setae appear to be pri-
marily bipectinate with occasional branches slightly out of line.
Often in mounting, however, flow of the medium causes all of the
branches to point in the same direction; also, the setae may become
twisted, so that the exact nature of the branching may be obscured.

On the genu, unipectinate setae are found on the dorsal margin
and bipectinate setae on the ventral margin, with intermediate
conditions on the lateral portions. The intermediates are more
stiffs than the ventral setae and are occasionally unipectinate.

Setae on the femur are similar to those of the genu, except that
the dorsal setae are usually bipectinate and differ from the ventral
setae only in being more stiff, although not as straight as the
dorsal genual setae.

The trochanter is a small, somewhat triangular segment. This
shape is fairly constant within species, and most of the species
have similar trochanters. Species of Cheladonta are characterized
by a noticeable elongation of the trochanter (fig. 85). Whartonia
senase (fig. 60) and Acomatacarus plumosus (fig. 88) have short
broad trochanters, otherwise no distinction among species was
noted. There are no setae on the trochanter. The articulation
between trochanter and femur does not appear to be flexible.

The femur is the longest palpal segment. The shape of the
femur is similar in all species. It is narrowed at the proximal end
and broadened at the distal end. In Neoschongastia (figs. 71-72)
the femur is extremely broadened. It is slightly arched and the
apparent shape varies somewhat with the position of the palpus
on the slide. A number of setae are borne on the femur and
taxonomic characters were found in the number and nature of
these setae. In general, the dorsal setae are thicker and stiffer
than the ventral ones. Elongated setae are often found on the

156 The University Science Bulletin

lateral and medial portions of the femur. Setae with reduced
branching and occasional nude setae were noted in Blankaartia
and in Speleocola tadaridae (fig. 86). Leeuwenhoekia americana
has distinctive dorsal femoral setae. In Acomatacarus arizonensis, A.
plumosus, and Whartonia senase some of the femoral setae are
expanded (fig. 88). The dorsal femoral setae of Trombicula
(Eutromhicida) belkini (fig. 79) have many more branches than
do those of other members of that subgenus.

The mean number of femoral setae for each species is given in
the descriptions of species. To obtain this figure the setae were
counted on both femora and an average number obtained for each
specimen. The means given are based on these average values.
Occasionally one femur is obscured to the extent that the setae
cannot be counted but usually, at least one femur is clearly visible
on each specimen. The number of setae on the femur is a char-
acteristic which may be used in separation of species. Usually,
related species have about the same number of femoral setae but
the exceptional cases provide taxonomic characters. Also, mem-
bers of the subfamily Leeuwenhoekiinae have a greater mean num-
ber of femoral setae than do the species in Trombiculinae ( see Table
3). No other categories are clearly separated.

It seems reasonable that the number of setae on a segment could
be correlated with size. Unfortunately measurements of palpal
segments are difficult to take accurately, due to distortion in mount-
ing, different angles of view, and lack of definite points from which
to measure. Body size also cannot be accurately measured on
mounted specimens. Therefore the length of the cheliceral base
was used as an index of size; the validity of this assumption is
admittedly open to question. The association between the number
of femoral setae and the length of the cheliceral base was tested
by Olmstead and Tukey's (1947) Corner Test for Association.
Within the species, the two were found to vary independently
in most cases. When the mean values for all of the species were
used, however, association was significant. Species with longer
cheliceral bases tend to have more femoral setae. It is inferred
that larger species have more femoral setae. That these two vary
independently within the species is also of interest.

The palpal genu is similar in shape in most of the species; it is
cylindrical, slightly smaller at the distal end, and about as long
or slightly longer than broad. The genu is wider than long in
species of Cheladonta, Pseudoschongastia, and Walchia americana.

Nymphs of the Trombiculidae

157

Some of the genual setae are expanded in Acomatacarus arizonen-
sis, A. plumosus, and Whartonia senase. Certain of the genual
setae are distinctive in Leeuwenhoekia americana. Dorsal genual
setae are occasionally nude in Speleocola tadaridae, Trombicula
crossleyi, T. fitchi, T. kardosi, T. merrihewi, T. montanensis, T.
ornata, and T. trisetica.

The number of setae on the genu was handled like that of the
femur, and means and standard errors are given in the descriptions
of species. No species groups are indicated by these figures but,
when considered with the mean number of femoral setae, an in-
teresting relationship emerges. The Leeuwenhoekiinae have a
greater mean number of setae on the femur than on the genu, while
the trombiculines have either a greater number on the genu than

GRAPH I: SETAE ON PALPAL FEMUR AND OENU

w
e

4S
40
86

80
25

ts
to

s •

SPECIES OF
TROMBICULINAE

SPECIES OF
LEEUWENHOEKIINAE

lb IS 20 25 30 35 40
MEAN NUMBER OF SETAE ON FEMUR

-I —
46

158

The University Science Bulletin

on the femur or about the same number on the two segments (see
Graph 1). This character holds fairly well on individual speci-
mens; of the trombiculines only four individuals were found which
had a greater number of setae on the femur, and in these cases the
diflFerence was one seta. The Leeuwenhoekiinae, as individuals,
have at least four more setae on the femur than on the genu.

As with the femoral setae, the number of genual setae was tested
for association with the length of the chehceral base. Again,

Table 3. — Mean Number of Setae on Palpal Femur:
Species among Taxonomic Categories.

Distribution of

Mean Number Setae

a)
o

3
(D

-2
"S

4)
O

"So
C

■§

3. . . .

5
2

1
1

■3'

1
1

2
1

within the species the two were found to vary independently but
between species, using average values, a positive association was
present (see Graph 2).

The palpal tibia is a small segment with a distal elongation bear-
ing the palpal claw and the accessory claws. The palpal tarsus
articulates to the ventral margin of the tibia. The distal elongation
is extreme in Blankaartia (figs. 169-172) and is suflBcient for identi-
fication of that genus. The palpal claw is slightly curved, never
divided as in the larvae, and usually about the length of the outer
tibial margin. Claws are difficult to compare as to shape, since on
the slide a slightly turned claw presents a difi^erent aspect. Within
a series of specimens gradations from an abruptly tapered claw
to an attenuated claw may be seen. The claw appears to be coni-

Nymphs of the Trombiculidae

159

Table 4. Mean Number of Setae on Palpal Genu:
cies among Taxonomic Categories.

Distribution of Spe-

Mean

Number

Setae

h-5

s
fe5

0
§

S
0

03
G

1
1

i
2

3
1

"■3"

1
1

"2"

1
1

1 ('D'>'r>'!xirplli\

^ \F

1 {Blanhaartia
velascoi)

1
1
1
1

cal or nearly so, and is always pointed. The claws of Neoschongas-
tia (figs. 100-101, 193-194) are considerably elongated, being more
than twice the length of the outer tibial margin. In Blankaartia
the claws are shorter than the tibia, which is elongated. The ac-
cessory claws are at the base of the palpal claw, on or near the
dorsal margin and they project medially and forward. The tips of
the accessory claws are spatulate;^ when seen in side view they
appear to be sharply pointed but are actually blunt. Most of the
species have two accessory claws in tandem, i. e., one behind the
other. In Bhnkcmrtm the number is variable, ranging from two
to seven, and the accessory claws are not all in tandem. In Neo-
schongastia americana the lower accessory claw is absent and a
nude or branched seta is present in the same position. In Neo-
schongastia hrennani the lower accessory claw is decidedly smaller
than the upper one. Otherwise the two accessory claws appear
to be about the same size and length though often they are not
parallel and are difficult to compare.

160

The Univ'ersity Science Bulletin

Both the claw and the accessory claws are probably modified
setae. There seems to be a gradient of stiffening setae on the
dorsum of the palpus which involves the straight femoral setae,
the monopectinate setae of the genu and tibia, the accessory claws,
and the claw. One specimen of Speleocola tadaridae had a
branched seta in place of the claw; the substitution of a seta for an
accessory claw in Neoschongastia americana has been mentioned.

3
Z
Ul
O

-i
<

z
o

es -

£4

80 -

18 -

2
Z

12 -
10 -

8 -
6
4
t

GRAPH 2: PALPAL SETAE IN RELATION TO SIZE •
MEAN NUMBER OF SETAE ON PALPAL GENU VS.
MEAN LENGTH OF CHELICERAL BASE.

+ • TROMBICULINE SPECIES

o • LEEUWENHOEKIINE SPECIES

+»

I +

+
+

MEDIAN

■ +

CORNER tests:

TOTAL - N= 46

-C • SO

LEEUWENHOEKIINE SPECIES
ONLY- N ' 8

.£?• IS**

TROMBICULINE SPECIES
ONLY - N • 38

>, • 19

— I 1 1 1 1 1 1 1 1 1 1 '

SO 60 TO 80 90 100 110 120 ISO 140 ISO 160 ITO ISO

MEAN LENGTH OF CHELICERAL BABE If)

The medial (inner) surface of the tibia usually bears one
branched seta adjacent to the articulation with the palpal tarsus.
This seta is lacking altogether in species of Neoschongastia and
Hannemania. In Blankaurtia alleei, Speleocola tadaridae, Trom-
bic.ula crossleyi, T. merrihewi, and T. trisetica the seta is often
nude. In other species the seta is occasionally duplicated or de-
leted. Additional setae on the medial surface are regularly found
in Neoschongastia americana, where a seta is substituted for the

Nymphs of the Trombiculidae 161

lower accessory claw, Chatia setosa, which has one to three addi-
tional setae, and species of Hannemania, which have an additional
seta below the accessory claws. This latter seta is very stiff and
unipectinate with fine branches. Setae on the dorsal margin of
the tibia have been counted with those of the lateral surface, al-
though occasionally one appears to be on the medial surface.

Several setae, usually four or five, are present on the lateral
(outer) surface of the tibia; these include an apical unbranched
seta, stiff unipectinate dorsal setae, and flexible ventral setae.
Usually the separation between types of dorsal and ventral setae
is not so distinct as on the genu. The apical nude seta is invariably
present and rarely with one or two fine branches.

The palpal tarsus is attached to the ventral margin of the tibia;
it forms the "thumb" of the chelate apex. The tarsi of the different
species have much the same shapes; a narrow, tapered segment,
rounded on the end. Tlie tarsus is subject to a large amount of
individual variation in shape, to which the mounting process prob-
ably contributes. The shape of the tarsus is not sufficiently re-
liable to characterize the species investigated. The tarsal setae,
however, provide several characters.

Typically the tarsus has about nine branched setae, about five
apical nude setae, and a striated tarsala. In those species with
nine branched setae, the branched setae are remarkably constant
in arrangement. On the dorsal margin is a row of three branched
setae decreasing in size distally. These setae are strongly uni-
pectinate, with thick shafts and fine branches. The other branched
setae have thinner shafts, fewer branches, and are less obviously
unipectinate or bipectinate. These are fairly constant in position;
two are found on the lateral surface, two on the medial surface,
and two on the ventral margin. In the drawings six of the branched
setae are included on the lateral smface and three on the medial
surface for purposes of comparison. Often it is not possible to
tell to which surface the dorsal or ventral setae might belong.
Variations in the number of branched setae on the tarsus are fre-
quent since there may be adidtional setae on the lateral surface
or lack of lateral or medial setae. Species of Hannemania and
Chatia setosa have up to eighteen branched setae. Euschongas-
toides, Pseudoschongastia, and Walchia americana have five to
seven branched setae on the tarsus. Species of Neoschongastia and
Speleocola tadaridae have tarsal setae nude or with reduced branch-
ing. In Neoschongastia such setae are short and stout.

6—8920

162 The Univ'ersity Science Bulletin

The tarsala is located on the lateral surface of the tarsus and is
probably invariably present. The position of this seta varies within
the species from near the base of the tarsus to about the midpoint.

Several short nude setae are present on the tip of the tarsus.
These are modified branched setae of the thick-walled type and
rarely, tiny branches may be seen. The number and arrangement
of these apical nude setae are sufficiently constant for characteriza-
tion of some species. It is usually possible to refer these setae
either to the inner or the outer surface. Those species with five
apical nude setae usually have three inner and two outer. Devia-
tions from five apical nude setae were found as follows. Six apical
nude setae (four inner, two outer): Leeuicenhoekia americana.
Four apical nude setae ( three inner, one outer ) : Euschongastia
criceticola, E. cynomtjicola, E. diversa, E. peromysci, E. pipistrelli,
Trombicula montanertsis, and T, myotis. Three apical nude setae
(two inner, one outer): Acomatacarus plumosus, Speleocola
tadaridae, Trombicula crossleyi, T. merrihetvi, T. ornata, and T.
trisetica. Two apical nude setae (one inner, one outer): species of
Euschongastoides, and Pseudoschdngosfia. One apical nude seta:
outer surface, species of Neoschongastia; inner surface, species of
Cheladonta and Walchia americana. Species of Hannemania have
five apical nude setae arranged four inner, one outer. Blankaartia
alleei has four apical nude setae, these being t\vo inner and two
outer.

Occasional specimens possess one apical nude seta more or less
than other members of the species. This variation was noted in
Trombicula whaiioni (five or six setae), Euschongastia jonesi and
E. setosa (four or five setae), and Chatia setosa (six or seven setae).

The Scutum

The scutum of trombiculid mites is a simple plate in the larvae,
but in the n) mphs and adults it is highly modified. The scutum is
probably the most frequently illustrated feature of postlarvae and
early descriptions of trombiculids are devoted largely to this struc-
ture. The scutum bears the sensilla or pseudostigmatic setae. The
exact function of these setae is unknown but they appear to be pri-
marily tactile organs. The modification of the scutum itself, espe-
cially the sensillary area, suggests that it has a particular function,
however, a thorough study of this structure has never been made.

Usually the scutum is considered to consist of three regions: An
anterior tectum, a rodlike crista, and a posterior sensillary area. The
tectum is a membranous flap extending from the anterior end of the

Nymphs of the Trombiculidae 163

crista above the gnathosoma and usually bearing one or two setae.
The crista is a heavily sclerotized rod lying on the surface of the
integument. Posteriorly, the crista expands into the sensillary area.
The sensillary area is a complicated structure, apparently hollow
internally, usually strengthened with ridges, and bearing the sensilla
at its sides. In the mounting process, the shape of the scutum is
usually somewhat distorted, particularly the sensillary area becomes
crushed. Also, the scutum but rarely turns on its side, so that a
dorsoventral view is most commonly seen. The exact nature of the
sensillary area is difficult to make out in this view, but for purposes
of taxonomy this position is preferable.

The combination of mounting distortion and dorsoventral ex-
pansion of the sensillary area increases the amount of variability
seen among mounted specimens. The general aspect of the sensil-
lary area among a series of similar specimens is often quite variable,
so that characterization of the scutum for a given species may be
difficult. Some previous attempts at separation of similar post-
larvae on scutal characters have failed due to this variability. When
a characterization is built upon a series of specimens some of
the features are found to be relatively constant, but these features
must be used with caution. An effort was made to use typical speci-
mens for illustrations, but to handle adequately the variation within
a species would require in some cases a drawing for every specimen.
In spite of this variation, some features are sufficiently constant as
to permit characterization of species and species groups.

The scutum is a relatively simple structure in the leeuwenhoekiine
species but becomes more complicated in the trombiculines. The
description of this structure may be handled profitably by beginning
with the simple types and noting the changes that occur in the more
complicated types. This is not intended to be a phyletic sequence
except in the most general sense. The elements of phylogeny are
certainly involved but the series of examples is not necessarily in
the proper order.

The scutum in the Leeuwenhoekiinae (figs. 138-143). These
species have scuta which appear intermediate in morphology be-
tween the trombiculine species and species of the family Trom-
bidiidae, particularly the subfamily Microtrombidiinac. They differ
from the latter in having two tectal setae rather than six or more.

The tectum of the Leeuwenhoekiinae is rather well sclerotized;
the margin of the tectum is smooth and never bears serrations or
teeth. The tectum is small in species of Hannemania, intermediate
in species of Acomatacarus, and large in Chatia setosa, Lee.uwen-

164 The University Science Bulletin

hoekm atnericana, and Whartonia senase. In the latter three species
the marghis of the tectum are heavily sclerotized and the shape of
the tectum is somewhat sagittate. In species of Trombidiidae
which have such a sagittate, projecting tectum, the term "nase" has
been applied to the structure. The tectum bears a pair of setae,
usually in the midregion but apically in Hannernania and Acomafa-
cariis. These setae are not always side by side and seemingly their
position may be disturbed by the mounting process. On the single
specimen of Whartonia senase these setae are almost in tandem.
The tectal setae bear short branches and differ from the body setae
if the latter are modified. In Acomatacarus plumostis (fig. 138)
the tectal setae resemble the body setae, being somewhat expanded.
In A. arizonensis (fig. 142) the tectal setae are not expanded al-
though the body setae are. The tectal setae of Whartonia senase
(fig. 143) are forked while in Leeuivenhoekia americana (fig. 140)
they are unusually elongated. The tectal setae are set in pits which
may have sclerotized margins. These are particularly evident in
Hannemania dunni.

The crista is a thin rod with somewhat irregular margins and is
readily subject to distortion in mounting. The connection of the
crista with the tectum is not always visible. It may be but lightly
sclerotized and is sometimes obscured by body setae. The crista
is rather broad in Acomatacarus arizonensis and Chatia setosa.
Some punctations may be visible on the crista and are prominent
in Chatia setosa (fig. 141). The crista appears to be on the sur-
face of the integument, since integumental striations never run
over it. Actually the crista may lie in a depressed area; at least
that of Chatia setosa gives this impression.

Beneath the sensillary area is a continuation of the crista, the
basal plate. In the leeuwenhoekiines this plate may be identi-
fied as two sclerotized bands, continuous with the crista ante-
riorly and uniting posteriorly to form a ventral apodeme, which
projects behind the sensillary area proper and is mostly subintegu-
mental. The posterior limits of the apodeme are sometimes hard to
find. In the illustrations the basal plate is indicated by dotted lines
on one side of the sensillary area. The sensillary area is usually
wider than the basal plate, but nearly coincides with it in Acomata-
carus arizonensis, Chatia setosa, and Leeuwenhoekia americana.
The space between the two bands of the basal plate hes beneath
the center of the sensillary area and is oval in Acomatacarus arizo-
nensis, and circular to triangular in other species.

Nymphs of the Tbombiculidae 165

The sensillary area proper is on the surface of the integument
and is continuous with the crista anteriorly. The center of the
sensillary area is a continuous plate which is usually broken in
mounting. It may be normally extended dorsally to form a dome-
shaped structure which is continuous with the body cavity through
the opening in the basal plate. The breaks which are seen on
almost all specimens, and which presumably occur during the
mounting process, are never regular and vary from specimen to
specimen. However, some species show a rather constant structure
which may indicate lines of weakness or actual sutures. In Hanne-
mania eltoni, an inverted triangle with ridges as borders is seen on
most specimens (fig. 139). Whatever the nature of the central
part of the sensillary area, the individual variation appears to rule
out taxonomic characters.

The shape of the sensillary area is roughly circular except in
Acomatacarus arizonensis, which has a transverse oval sensillary
area. The area is bordered by ridges posteriorly and at the sides;
these are irregular in most specimens and are probably also dis-
torted in mounting.

The sensillary bases or pseudostigmata are found at the sides of
the sensillary area to midway between the sides and the middle of
the area (Acomatacarus arizonensis, Chatia setosa). These appear
to be conical pits. Around the base of the sensillum itself is a
heavily sclerotized ring, and at the outer margin of the pit is a
similar larger ring. Between these two are several less distinct
concentric circles. Jones (1950:487) has described the sensillum
and sensillary bases of the larva of Tromhictila atitumnalis. Their
structure is similar (o that of the nymphs and suggests their func-
tion to be "a sensory perception of touch and probably of vibra-
tions in the air."

The sensilla in the Leeuwcnhoekiinae are flagelliform and un-
branched. There appear to be no tiny basal branches. The length
of the sensilla is not readily measured because the structures are
rarely sufficiently straight and because their very fine tips disappear
among the body setae. No characters associated with the sensilla
were found among species of Leeuwcnhoekiinae.

A possible character mentioned by Audy (1954:131) involves
"parascutal" setae. These are the body setae which are found at
the sides of the crista in front of the sensillary area. Since these
setae merge with the other body setae behind the sensillary area,
it is not possible to enumerate them accurately in the Leeuwen-

166 The University Science Bulletin

hoekiinae. In Hannemania there are about fifty such setae, and
in smaller species (Acomatacarus arizonensis) about ten or fifteen.
These setae are morphologically similar to other body setae, the
most anterior of them being slightly smaller. In the illustrations,
setal bases beside the crista are indicated and one body seta is
figured. Other setae occur lateral to these in the prescutal area.
Puncta are seen occasionally in the sensillary area. They are
found in the anterior area in Chatia setosa and Leeuwenhoekia
americana and scattered over the sensillary area in Hannemania;
other species may show scattered puncta on some specimens.

The scutum in the Trombiculinae (figs. 138-168, 173, 180). In
general, scuta of trombiculines differ from those of leeuwenhoekiines
in having one or no tectal setae instead of two, in having lateral
keels along the crista (in most cases), in having branches on the
sensilla (usually), and in having a sensillary area which is struc-
turally more complicated.

Modifications of the scutum in the trombiculine genera allow
three types of scuta to be recognized. These are discussed sep-
arately below. These types tend to grade into one another but the
division is of use in the discussion.

"Type A" is possessed by Euschongastoidcs hopJai, E. Joomisi,
Pseiidoschongastia farneri, P. hu7igerfordi, and Walchia americana
(figs. 144-147). These species have a scutum with a rather short
crista and a broad sensillary area which is less complicated than
the following two types.

The tectum is broad and short in all but Walchia americana,
which has an almost square tectum. Species of Pseiidoschongastia
lack serrations on the tectimi. Small serrations are present on
the anterior border of the tectimi in species of Euschongastoidcs,
and large, long serrations are found on the teclum of Walchia
americana. The tectal seta is lacking in Walchia americana and
is very short in the other species. Branching is much reduced in
all except Euschongastoidcs hoplai. The connection of the crista
with the tectum, and the posterior portions and corners of the
tectum, are diflScult to see in these species, except for Walchia
americana.

The crista is a rather narrow, heavilv sclerotized rod. Lateral
keels (Brown, 1952:20) appear to be present in all species. These
are flaplike, membranous structures lying beside the crista; they
are invariably displaced dming the mounting process so that their
exact position is not known. Apparently they parallel the crista on

Nymphs of the Trombiculidae 167

each side and connect posteriorly with the sides of the bulla (see
below). Lateral keels were not visible in all specimens, and usually
only portions of them could be seen. They are most clearly visible
in Walchia americana.

The crista joins the sensillary area and immediately divides into
two arms which run laterally at right angles to the crista. These
form the anterior margin of the basal plate. Among scuta of this
type the basal plate consists of two halves lying approximately
under the sensillary bases. In Euschongastoides the plates also
appear to have a thin posterior connection. The basal plate con-
tains openings for the sensillary bases. The shape of the basal
plate is slightly different among the three genera, the anterior
parts being somewhat difficult to see. At the anterior portion of the
sensillary area a somewhat triangular structure occurs, which has
its apex on the posterior part of the crista and its base in the middle
of the sensillary area. The term "bulla" seems appropriate for
this structure, since it seems to be hollow and domelike, and ex-
panded dorsally. The bulla is continuous with the crista anteriorly;
small ridges or bars connect the corners to the sides of the sensil-
lary area; and the lateral keels appear to be attached at these cor-
ners. The posterior margin is convex and is usually irregular.
The sensillary bases are similar to those of the leeuwenhoekiine
species; they are located at the lateral extremes of the sensillary
area. Ridges encircle the sensillary area, running from the sides
of the bulla around the sensillary bases and along the posterior
margin of the area. These ridges are irregular in shape and are often
broken and displaced in mounting. In Walchia americana the
major ridges pass medial to the sensillary bases instead of around
them; small ridges are found lateral to the sensillary bases. Also,
in Walchia americana the posterior ridge is discontinuous and con-
nected by a small secondary ridge at the midpoint of the posterior
end of the area. Body striae enter the sensillary area over the
anterior ridge, passing between the bulla and the sensillary bases;
the striae are continuous through the depressed area behind the
bulla but do not cross the posterior ridge. The posterior apodeme
was clearly seen only in Walchia americana, where it is much re-
duced.

All of these genera have sensilla with branches. In species of
Euschongastoides the sensilla have very minute branches on their
extreme basal portions; these are followed by longer branches for
about half the length of the sensillum. The distal half is provided

168 The Unintrsity Science Bulletin

with much longer attenuate branches. The stem of the sensillum is
long and attenuated; near its apex it becomes zigzag in most speci-
mens (fig. 146). Branches arise from at least two planes of the
sensillum. The sensillum of Wolchia amcricana is similar to that of
Eiisclwngostoides but the stem is somewhat thicker and straighter.
The sensillum of Fseiidoschdngastia farneri has a rather thick
straight stem, with short basal branches, long attenuate branches
in the midregion, and shorter attenuate branches at the apex (fig.
145). The medial branches have enlarged bases, giving the stem
a slightly irregular appearance. The sensillum of P. hungcrfordi is
similar to that of F. farneri but is usually somewhat expanded medi-
ally, presenting a subclavate appearance (fig. 144). Occasional
specimens of P. hungcrfordi have the sensillum very little expanded
or not at all, so that it alone would not serve to separate the two
species of Pseudoschongastia. One series of P. ]iii7igerfordi con-
tained no specimens with expanded sensilla.

Scutal punctations are occasionally present but never numerous in
the sensillary area or on the crista.

The parascutal setae are much less numerous than in the leeuwen-
hoekiine species. Usually they appear to be relatively distinct from
the other body setae due to their decreased density. Precise
enumeration is difficult; six to fourteen parascutal setae were noted
among these species, except for Wolchia amcricana. This latter
species consistently has but a single pair of parascutal setae. This
appears to be a sound taxonomic character as no specimens of other
species were found with but two parascutal setae.

The two remaining types of scuta are similar to each other.
"Type B" includes species with sensillary bases rather far forward
in the sensillary area and the central portion of this area reduced,
so that the sensillary area has a reniform shape. "Type C" includes
species with the sensillary bases at the middle or back of the sensil-
lary area with the central portion of this area modified and promi-
nent, the appearance of the sensillary area being triangular or trans-
verse. The distinction between these two types, while not clear-
cut, is one of convenience.

Species possessing "Type B" scuta are Cheladonta michcneri,
C. ouachitcnsis, Neoschongastia amcricana, and N. hrennani (figs.
149, 154-155, 180). In these species the tectum is provided with
serrations or teeth; these are long in species of Neoschongastia and
short and broad in species of Cheladonta. The tectal seta is longer
than the body setae.

Nymphs of the Trombiculidae 169

The crista is rodlike and plainly connected to the tectum.
Lateral keels are present although distorted and often difficult to
see. The crista broadens before it reaches the sensillary area in
Cheladonta; in Neoschongastia the crista reaches the sensillary area
before broadening.

The basal plate of the sensillary area in these species arises as a
furcation of the crista. In Cheladonta (fig. 154) it consists of tsvo
bands passing posteriorly beneath the sensillary bases; an opening
in the basal plate is found beneath each sensillary base. The basal
plate of Neoschongastia (fig. 180) is similar but has a larger, drop-
shaped opening for each sensillary base. In Neoschongastia the
basal plate appears to terminate at the posterior margin of the
sensillary area; the apodeme is either much reduced or absent. The
basal plate of Cheladonta, however, plainly continues posteriorly to
the sensillary area as an apodeme formed from the union of two
bands. The apodeme is broad, often marked with irregular openings
and often furcate posteriorly.

The bulla of the sensillary area is somewhat reduced in size in
Cheladonta but still triangular in shape; in Neoschongastia the bulla
is much reduced and is hardly wider than the crista in N. ameri-
cana. Ridges from the corners of the bulla pass later ad to the
sensillary bases and unite at the back of the sensillary area. In
NeoS'Chongastia additional ridges pass medial to the sensillary
bases. These ridges are variable in shape and position, apparently
due to distortion in mounting. Body striae enter the depressed
area behind the bulla from the anterior angles of the sensillary area.
Body striae also surround the sensillary bases but do not cross the
posterior ridge of the sensillary area.

Punctations are present on the posterior part of the crista and
in the sensillary area.

The sensilla are provided with branches, which are tiny on the
basal portions and become longer distally. In Cheladonta mich-
eneri they are subclavate in shape; in C, otiachitensis they are
slightly expanded in their midregions but taper again in the distal
portion. The sensilla of Neoschongastia americana are similar but
slightly expanded distally, and with short branches on the basal half.

Parascutal setae are similar to the body setae but less dense,
numbering about ten.

A "Type C" scutum (sensillary area triangular rather than reni-
form) characterizes the genera Blankaartia, Euschongastia, Speleo-
cola, and Trombicida (figs. 4, 148, 150-153, 156-168, 173). These

170 The Univtsrsity Science Bulletin

genera include about one half of the species in this study. The
"Type C" designation is essentially a category for those scuta with-
out unusual modifications.

The tectum is square or triangular in most species; the margin
of the tectum is smooth in Blankaartia and Euschongastia but pro-
vided with serrations or teeth in Speleocola and Trombiciila. In
most species the teeth number about eight to twelve; often they
are difficult to see and counts seem to be impractical but the pres-
ence or absence of teeth usually can be ascertained without diffi-
culty. In the subgenus Etitromhicttla the teeth are smaller and
more numerous, averaging about twenty. A slight central prolon-
gation of the tectum is present in species of the subgenus Neotrom-
bicula and in Trombiciila ornata. A single tectal seta is present in
all species; occasionally it is duplicated and in these cases the two
setae are in tandem. The tectal seta tends to be elongated in most
species of Euschongastia.

The crista is a narrow rod, often of irregular width. Lateral keels
were seen in all species except Blankaartia velascoi; these are in-
variably disturbed in mounting and their terminations are obscured.
The sensillary area may be roughly characterized as a transverse
oval in most Euschongastia, compared to a triangular shape in
the other species. The basal plate is composed of two halves,
formed by a furcation of the crista anteriorly; the posterior termi-
nation is difficult to see but apparently the two halves are united
beneath the posterior ridge. The plate contains openings for the
sensillary bases ( these are omitted from the illustrations ) . As with
the previous species, the basal plate is indicated on one side of
the illustrations by dotted lines. The shape of the central opening
of the basal plate may be a useful character for separating some
species. The apodeme may be seen as an extension of the basal
plate in most Trombiciila but is reduced to a tiny isolated piece in
T. crossleyi, T. 7nerrihcwi, T. ornata, T. trisetica, and species of
Euschongastia except setosa.

The bulla is present in all species and is essentially the same
shape in most species; an expanded tiiangular structure at the
anterior portion of the sensillary area. The bulla is reduced in
size in Euschongastia diversa, E. peromysci, Speleocola fadaridae,
and Trombicula myotis; the bulla is elongated anteriorly in species
of Blankaartia. A median carina is present atop the bulla. This
structure is usually flattened in mounting and may be obscured
or confused with the walls of the bulla; apparently all of the species

Nymphs of the Trombiculidae 171

have a carina but it is not always seen. The carina is thickened in
some species, particularly in the subgenus Neotromhictila. In this
group the carina appears to be a thick, slightly elevated structure
but may actually be absent. In occasional specimens of many
species the carina is doubled at the posterior end.

The sensillary bases have the same structure as described for
the other types of scuta. The bases are placed well at the sides;
they appear slightly enlarged in TromhicuJa crossleyi, T. montanen-
sis, T. merrihewi, and T. trisetica. This may be a function of the
small size of these scuta.

The sensillary area is surrounded by ridges, which include weak
ridges from the angles of the bulla, heavy ridges around the out-
sides of the sensillary bases, and a rather weak posterior ridge.
The ridges encircling the sensillary bases have elevations in front
of and behind the bases. A crossbar may usually be found medial
to the sensillary base but is usually weak and often distorted. In
Speleocola todaridae these crossbars are strong and have elevated
flanges on them (fig. 173). Additional crossbars are found irreg-
ularly on some specimens. The posterior ridge is usually broken in
mounting. In species of the subgenus Eutrombicida this ridge
has a bilobed appearance caused by two elevated areas, a con-
dition indicated in TromhicuJa gurneiji. Body striae enter the
sensillary area at the front, between the bulla and the sensillary
bases, while a few striae appear to cross the ridges around the sen-
sillary bases. Striae do not cross the posterior ridge.

On the margin of the sensillary area, near the angles of the bulla,
one or two tiny projections may be seen. These are apparently
subcutaneous and are inconspicuous except in the genus Euschon-
gastio. In species of Euschdngastia (except E. trigenuala) these
projections are enlarged into rather conspicuous teeth. The limits
and extent of the teeth are difficult to make out, owing to confusion
with body striae and with lateral keels. Their exact nature is not
known.

The sensilla afford usable characters m this group of species.
In species of Blankaartia the sensilla are long, attenuated, and
entirely nude. Sensilla of other species have, at least, tiny basal
branches. Species having sensilla which are rather thick but not
subclavate and provided with short branches along the entire
length include Euschongastia criceticola, E. cijnomyicola, E. di-
versa, E. peromysci, E. pipistrelli, E. setosa, TromhicuJa autum-
nMs, T. fitchi, T. JipovsJcyi, and T. syJviJagi. Sensilla of this type

172 The University Science Bulletin

but with a reduced number of branches are found on Etischongastia
ionesi and Trombicula kardosi. The sensillum of Eiischongastia
trigenuala is similar but lacks branches on the distal third of its
length. The sensillum of Trombicula tvhartoni is similar in being
rather thick but not subclavate, but lacks branches except for
minute basal ones. Species having sensilla slightly expanded dis-
tally, or subclavate, and provided with rather heavy branches
include Speleocola tadaridae, Trombicula crosslcyi, T. merrihewi,
T. ornata, and T. trisetica. Sensilla which are flagelliform, having
thin stems, and provided with long fine branches are possessed by
Trombicula belkini, T. gurncyi, and T. hansaseiisis; similar sensilla
with numerous fine branches are found on T. montanensis and T.
myotis. Species with similar sensilla but which have the branches
less numerous and arising from a single plane include members
of the subgenus Eutrombicida, except T. belkini. In the subgenus
Neotrombicula, many specimens have sensilla with furcations at
the tip; these are rather small and appear to be two normal branches
rather than an additional modification of the stem.

A few puncta may be seen in the sensillary area on most speci-
mens. Puncta are exceptionally numerous on species of the sub-
genus Eutrombicula (except T. belkini), Trombicula lipovskyi, and
T. tvhartoni.

Parascutal setae were found to be difficult to count precisely;
most species have about eight to fourteen such setae except for
members of the subgenus Eutrombicula, which have about twenty
to thirty, and species of Blankaartia, which have about thirty. Ex-
cept for these extreme cases, the number of parascutal setae would
be a difficult character to apply. In Trombicula crossleyi and T.
trisetica a pair of parascutal setae is placed directly in front of the
sensillary area, one seta on either side of the crista. These setae
are noticeably longer than the other parascutal setae. Trombicula
ornata has two parascutal setae in the same position which are but
slightly longer than the other parascutal setae. In other species,
there are usually no setae in this position.

Measurements of the scutum. A series of measurements of the
larval scutum has been used successfully for some years. Workers
on postlarval stages have taken various measurements of the scutum
but there has been no general agreement as to what to measure or
how the measurements are to be taken. Audy (1954:128) has pro-
posed an ambitious scheme for taking eleven measurements of the
scutum; his system includes the better ideas of other workers but is

Nymphs of the Trombiculidae 173

still affected by an underlying difficulty, in that the nymphal scutum
has few definite points from which to originate or complete a meas-
urement. The larval scutum, in contrast, has five or six scutal seta
in addition to the sensilla, and the larval measurements are based
upon these definite points.

Both the anterior and the posterior margins of the sensillary area
are indefinite; the anterior margin is marked by the gradual widen-
ing of the crista, and the posterior margin by the very irregular
posterior ridge. Thus it seems that measurements of the length of
the crista itself, the length of tlie sensillary area, the length of the
posterior apodeme, or others involving these points would prove
impractical. The writer has taken only four measurements, as fol-
lows: ASL: Anterior scutal length, measured from the tectal seta
to the level of the centers of the sensillary bases. SB: Distance
between sensillary bases, measured from the point of insertion of
the sensilla. TS: Length of the tectal seta(e). SENS: Length of
the sensillum. The ratio ASL/SB provides a quantitative expres-
sion of the general scutal shape.

The debatable measurement here seems to be the measurement
ASL. The anterior point used could have been the anterior termi-
nation of the crista rather than the tectal seta. However in many
species the measurement cannot be taken with the oil immersion
lens, and the tectal seta is visible under lower magnification al-
though the anterior termination of the crista may not be. At lower
magnification it is easier to determine the posterior limit of the
measurement; the ocular scale used by the author had long rulings
which often spanned the distance between the two sensillary bases.
A further difficulty arises in the genus Walchia, which lacks a tectal
seta. For the single species included in this work, the measurement
was taken to about the middle of the tectum ( a procedure suggested
by the position of the tectal seta in Pseudoschongastia) . Workers
in the Asiatic Pacific area have to^ contend with many species of
Walchia and may well find this procedure objectionable.

Accurate measurements of the lengths of the sensilla are impos-
sible except in the species where they are thickened and straight.
In other species estimates of the length were made.

The measurements ASL, SB, and TS are given in the descriptions
of species. The measurement ASL is roughly related to the size of
the mite and has a large standard error. The measurements SB and
TS are rather more independent of size and have smaller standard
errors. However, all three of these measurements mav be used to
separate some species from others, and as such, they provide usable
taxonomic characters.

174

The University Science Bulletin

The ratio ASL/SB is also given in the descriptions of species.
In table 5 the distribution of this ratio among taxonomic categories
is given. This table shows that three general groups are formed:
A lower group with mean ratios between 1.2 and 1.7, a central
group with mean ratios between about 1.9 and 3.0, and an upper
group with mean ratios from about 3.2 to 5.2. The upper group
is composed of leeuwenhoekiine species, but members of the genus
Blankaartia approach them. The lower group is composed of
Eiischongastoides, Pseudoschongastia, and Walchia (species with
a "Type A" scutum). Most species fall into the middle grouping.
It is further seen that the generic groupings are fairly distinctive.
As more species become known, it may develop that the ratio
ASL/SB is a valuable generic character.

The Body

This section considers the body and its morphological features
not previously treated, namely the setae, eyes, sternum, genitalia,
and anus.

Shape. Body shape has been frequently used to characterize
trombiculid postlarvae. A classic method for separating trombicu-
lids from their relatives was by body shape; trombiculids were be-

Table 5. The Ratio ASL/SB: Distribution of Species among Taxonomic
Categories. ( ASL and SB are explained in text. )

(Si

§.

•«.i
o

■*.o

Mean

■•o

Ratio

•5

.s a

fe;

fcq

1 2

1 4

1 (Walchia)

1 6

1 8

2 0

1
2
1
4

2 2

4
2
4

2 4

1
2
1

2 6

2 8

3.0

1 (B. alleei)

3 2

1 (/

I . arizovfm.-'iift)

1 {B. velascoi)

3.4

3 6

1
1

2
2

3.8

4.2

4.5

5.2

Nymphs of the Trombiculidae 175

lieved to have a constriction of the body which produced a "figure
8-shape," while trombidiids lacked such a constriction. Subse-
quently it was discovered that leeuwenhoekiine genera (except
Hannemania) lack the constriction; however, all known trombicu-
line species are constricted. This feature is often destroyed in
mounted material and must be used with caution except in fresh or
alcoholic material. Another difficulty in applying the body constric-
tion as a diagnostic character separating trombiculids from trom-
bidiids is the presence of slight constrictions in some trombidiid
species. The body shape of leeuwenhoekiines (except Hanne-
mania) is a slightly elongated oval (fig. 205). Most trombiculines
have a rounded "figure 8-shape" ( fig. 202 ) except for certain species
which are noticeably elongated; these are species of Cheladonta
(fig. 200), Pscudoschongastia, Trombicula aiitunmalis, T. crossleyi,
T. tnerrihewi, T. sylvilagi, and T. tmetica (fig. 1). An elongated
body has been suggested as a possible character for Neotrombiciila
but this is not a feature of all members of that subgenus. The body
is shortened and stronglv constricted in species of Neoschongastin
(fig. 201).

Size. As previously mentioned, body size of nymphs is not sub-
ject to exact measurement due to extreme distortion in mounting.
This is less true of the smaller species than the larger ones; never-
theless, most specimens could not be measured with any feeling of
accuracy. Estimates of the body length were made only to tenths
of a millimeter; these are given in the descriptions. The range of
sizes found was from 0.4 to 1.2 mm. Most of the species are be-
tween 0.6 and 0.8 mm. long; species measuring 0.5 or less were
characterized as "small" (seven species); those measuring 0.9 or
larger were characterized as "large" (nine species). No estimates
of width were made.

Eyes. The presence or absence of eyes, and their number and
position, is a much-used character in mite taxonomy. Eyes of
trombiculids are bright red in life but are sometimes difficult to see
in mounted mateiial. Most species lack eyes. Of the leeuwen-
hoekiinae species, all have eyes except Acomafacariis arizonensis
and Wlmrtonia senase. The eyes are found in front of the sensillary
area at the sides of the propodosoma or nearer the crista. The eyes
are double (2/2) in species of Hannemania but are single (1/1)
in other species. Of the trombiculine genera, eyes are found only
in the genus Blankaartia where thev^ are single and placed as in the
leeuwenhoekiine species, and in the subgenus Eutrombicula where
they are single and placed adjacent to the sides of the sensillary

176 The University Science Bulletin

area. In Blankaartia the eyes are distinct ovals; in Eutrombicidu
they are less distinct and sometimes have a semilunar appearance,
owing to their proximity to the sensillary area. The eyes are clearly
visible in T. alfredduge^i and T. splemlens, but hard to see in T.
belkini and T. lipovskyana. Particularly in T. beJkini, the presence
of eyes is detected in mounted specimens only by the disruption of
the cuticular striae. A lens could be seen clearly only in one speci-
men of belkini examined in this study.

Observations on living material in culture disclosed no obvious
differences in behavior between species lacking eyes and species
possessing them. Species lacking the lenses may well have light-
sensitive areas which are not detected in cleared and mounted
material.

Setae. A dense coat of body setae is characteristic of the group
of mites to which trombiculids belong. Characters involving body
setae have been very successfully used among these mites. Typi-
cally body setae are branched and so numerous as to form a thick
pelage which obscures the integument and even features of the
setae themselves. The dorsal setae are somewhat longer and
thicker than the ventral ones; those of both surfaces increase in
length posteriorly, so that the longest are the posterior dorsal ones.
Posterior setae typically arise from tubular elevations each of which
is supported by a small circular platelet. The ventral and the an-
terior dorsal setae arise from less distinct platelets and are but
slightly elevated.

Body setae are best observed at the posterior margin of the body.
Posterior dorsal setae have been figured for most species. In the
following discussions distinct types will be mentioned but not de-
scribed in detail, since the illustrations better fulfill this purpose.

Primitive body setae are probably the types with simple attenu-
ated stems and simple branches; however, of the leeuwenhoekiine
species only Chatia setosa has such setae. Among the trombiculines
most species have unmodified attenuated body setae although the
stem may be shortened or lengthened, and the tip of the stem may
be modified. Body setae with straight but thickened stems are
found in Hanncmania and some species of Trombicida.

A usual type of tip found on the posterior body setae is a division
of the attenuated stem into two or three short branches. Modifica-
tions of the tips are often so constant as to form valuable generic
or specific characters. Chotia setosa and Euschdng,astia pipistrelli
possess body setae with single, very long, attenuated tips. Species
of Har^nemania have body setae which terminate in a curved, al-

Nymphs of the Trombiculidae 177

most hooklike spine. Body setae of Trombicula mijoiis terminate in
hyaline knobs. Very tiny branches are found in groups at the tips
of die body setae of some species, including Euschongastoides
loomisi, Walchia americana, some species of Trombicula, and par-
ticularly some species of Etischongastia. In the latter genus a tran-
sition may be seen from a seta with a simple tip (E. criceticola)
through one with short branches on the tip (£. diversa, E. setosa)
and a seta with the stem and its tiny branches offset at the tip (£.
jonesi) to a seta with many tiny branches over the tip and even
distributed along a dominant lateral branch (E. peromysci) . Rela-
tively small branches are found near the tips of setae of most species
of Tromhicula.

Modifications of the usual shape of the setae include the pecul-
iarly expanded setae of Acomatacarus and Cheladonta, and the
leaflike setae of Whartonia senase. Compound branching is found
in the setae of species of Neoschongastia and Speleocola tadaridae.
The setae of Tromhicula fitchi are peculiar in the arrangement of
the branches; long branches are found on the basal half but only
short, thick-based branches occur on the distal half. Setae of T.
kordosi are similar to these but occasional long branches are found
among the shorter ones.

The increase in length posteriorly is usually a gradual transi-
tion, but in Euschongastia pipistrelli, E. trigentiala, and Trombicula
fitchi the change is abrupt; posterior setae are strikingly longer
than the adjacent dorsals. Intermixed long and short body setae
are found on Acomatacarus arizonensis and species of Neoschon-
gastia.

The lengths of various body setae have been used as specific
characters with some success. Certain difficulties arise, however,
when measurements of these setae are attempted. Modifications
of the tip often make the exact termination of the seta obscure;
lateral branches may exceed the tip of the stem in length and these
are so flexible that only an approximate length can be obtained.
Setae which end in fine tips cannot be measured accurately. Even
the relatively unmodified setae can be exasperating in this respect.
The writer found it impossible to obtain the same figures in re-
measuring setae of Trombicula alfreddugesi and T. splcndens.
Authors have recommended the measurement of setae from various
parts of the body, but since the setae increase in size posteriorly,
these measurements are useless unless the exact location of the
seta is specified. The writer measured only posterior body setae;
these figures are given in the descriptions as approximations. A

178 The University Science Bulletin

continuous range of values was found from about twelve to about
one hundred and five microns. Variation among individual speci-
mens seems to be about twenty microns in the species with larger
setae and less in those with smaller ones. The central posterior
dorsal setae on a single specimen appear to show less variation than
occurs between individuals. All in all the length of the posterior
body setae is a usable character if not too strictly applied; species
towards the extremes may be separated easily from others but spe-
cies with setae of similar length should be compared with caution
as to this character.

Sternum. The sternal area lies between the coxae of legs I and
II. Recently Womersley (1952:18) found several taxonomic char-
acters associated with the sternum. Some species (Giinfherana)
have precoxal plates of coxae I present, other species ( Trombicula,
subgenus Leptotromhidiiim) have these precoxal plates of coxae
I fused to form a longitudinally divided sternum, and most species
(most Trombictila) have the longitudinal division obliterated so
that a sternal area remains which is closed behind by a crossbar
connecting coxae II. In Gahrliepia (== Walchia) Womersley found
no evidence of a closed sternum formed by fusion of precoxal plates.

Of the species in the present work, only Neoschongastia was
found to have precoxal plates (fig. 177). (Womersley, 1952, in-
cluded descriptions of three species of Neoschongastia but did not
mention the sternum. ) A longitudinally divided sternum is present
in Trombictila (Leptotrombidium) myotis, which agrees with Wom-
ersley's findings.

The sternum in the leeuwenhoekiine species consists of a scle-
rotized plate bearing eight or ten setae. In the other ( trombiculine )
species the sternum is closed behind by a bar connecting the coxae
of legs II, with certain exceptions. Speleocola fodaridae, Trom-
bicida crossleiji, T. merriliewi (fig. 181), T. ornata, and T. trisetica
have sternal areas which are elongated posteriorly and open be-
hind; however, some specimens show a very weak incomplete bar
between the coxae II. Apparently the closure has been secondarily
lost in these species, perhaps as a consequence of change in shape
of the sternal area or decrease in general size. Other species have
a sternum of pentagonal to rectangular shape (or elongated in
Euschongastia jonesi and Walchia americana) which is closed be-
hind, although the posterior bar is weak in some species and some-
times is incomplete. (Walchia americana, in contrast to Womers-

Nymphs of the Trombiculidae 179

ley's observations on that genus, has a closed sternum.) Oc-
casional specimens (particularly specimens of Cheladonta and
Walchia americana) show a trace of a longitudinal division of the
sternal area. This would seem to lend support to Womersley's
suggestion that the closed sternum is derived from fused precoxal
plates. However, some leeuwenhoekiine species have a posterior
thickening of the sternal plate which resembles the closure of the
trombiculine sternum.

Setae on the sternum are usually of the ventral body type (but
different in Acomatacortis and Whartonia senase). About eight
setae are usually found in the sternal area; larger species have
about twelve setae and smaller ones about four to six. The setae
were not counted on all specimens but the variation noted was
large (about six to fourteen sternal setae in Trombiciila splendens).

Genitalia. The genital opening (fig. 12) is located between or
immediately behind coxae IV. The genital areas of all species
resemble one another very closely. Most external is a pair of
elongated plates which bear branched setae similar to those of
the body; the genital opening is between these two plates. Mesal
to these, and extending across the genital opening, is a pair of
elongated plates which bear the three pairs of genital setae. These
setae (figs. 13-15) are invariably six in number and are usually
nude but occasionally forked or branched. Rarely one or two of
the genital setae resemble the fanlike genital setae of the adult
male (see fig. 15). This phenomenon may be the expression of
an adult character in the nymphal stage, but it would appear that
sex is not consistently expressed in the nymphs in this manner
since very few specimens show this character. The two pairs of
genital suckers are located beneath the plates in the midregion
of the genital area. Their function is unknown. Their size ap-
pears to be variable, and their relative size was found to be in-
constant in a series of specimens.

Anus. The anus is located posterior to the genital opening. The
anal opening itself is not evident; the term "anus" is applied to the
plate or plates guarding the presumed position of the opening. In
the leeuwenhoekiine species (fig. 197) the single anal plate is cir-
cular or oval, poorly sclerotized, not obviously divided, and usually
distorted in mounting. The anal opening of trombiculine species
(fig. 199) is obviously guarded by two elongated shoelike plates,
each of which bears a number of setae. No further characters asso-
ciated with the anus were found.

180 The University Science Bulletin

Legs

Characters involving the legs of trombiculid postlarvae have been
virtually ignored, probably because of the plethora of setae found
on them. In the classification based on larvae, features valuable in
taxonomy include the number of leg segments and the number of
modified setae on certain segments.

All of the species included in the present study have, as nymphs,
seven segments in the legs; these are coxa, trochanter, basifemur,
telofemur, genu, tibia, and tarsus. A pretarsus, consisting of a pair
of claws, is present on the tarsus of each leg. Except for several
exceptions the relative shapes of the various segments appear to
be the same for all species. Leg setation, also, is very similar
among the various species. Therefore the following discussion will
be general, particularly in regard to leg setae.

Usually the legs are slightly shorter than the body, leg I being
the longest and largest. In Chatia setosa (fig. 190) the legs are
noticeably elongated, leg I being obviously longer than the body
and the other legs about as long as the body. Legs appear slightly
shortened in some species and particularly in species of Chela-
donta, but others (Euschongastoides, Neoschongastia, Pseudoschon-
gastiu, and some T romhicula) approach this condition. The coxa
is a rather flat, cylindrical segment which appears to be slightly
movable. The trochanter is a small, somewhat arched segment.
Articulations between coxa and trochanter and between trochanter
and basifemur are dicondylic; the former permits dorsal-ventral
movement and the latter anterior-posterior movement of the more
distal segments. Articulations between remaining segments are
monocondylic and dorsal. The distal segments are cyhndrical,
about as long or longer than broad. Shape of tarsus I appears dis- I
tinctive for certain species, these being the somewhat pyriform -3i
tarsi of Acomatacariis and Whartonia senase (figs. 187, 189), and 'i

the rather rounded tarsus of Cheladonta (fig. 184). Otherwise {j
tarsi are generally similar in shape. Walchia americana possesses I

a short stump which projects from the distal dorsal portion of tarsus
I (fig. 188).

Two measurements were taken on tarsi I, these being the
length (TL), measured from the dorsal articulation to the ventral
tip, and the thickness (TH), measured at the widest point. The
ratio TL/TH was computed. These figures with their standard
errors are given in the descriptions. Generally the size of the
measurement reflects the size of the mite. The ratio TL/TH has

Nymphs of the Trombiculidae 181

a rather small range among most species; however, the standard
error is small and this character is useful in separation of some
species. No groups of species were distinguished either by the
measurements or by the ratio.

Leg segments, especially the more distal ones, are covered with
setae. While the number of nude or branched setae on a segment
is constant for the species in many larvae, the number of boih types
varies among specimens of postlarvae. Audy (1954:128) has pro-
posed a scheme for tabulating the number, type, and location of
leg setae. The writer used a somewhat simplified modification but
abandoned the project as being laborious and unproductive. All
in all, the tabulation was completed for six species; the following
description is based upon this information and upon less precise
observations on the other species.

Interspecific variation in the number of setae per segment was
found in all species and on all segments. Coxal setae resemble
the ventral body setae; straplike extensions of the coxae I extend
towards the dorsum and these bear elongated setae. The trochanter
bears branched setae which are elongated and placed in a band
across the segment; the number of setae found was three to five
on the smaller species and seven to nine on the larger ones.
Branched setae on the remaining segments become increasingly
modified distally; the setae become unipectinate and the branches
become straight and increasingly fine. Setae on the tarsus may be
described as comblike. Branched setae on the legs are slightly
expanded in species of Acomatacarus, Leeuwenhoekia americana,
and Whartonia senase. Nude setae first appear on the telofemur,
which has zero to two dorsal nude setae. The remaining distal
segments have increasing numbers of nude setae, particularly on
leg I. On tarsus I nude setae are extremely numerous. On the
remaining leg segments the nude setae do not become so numerous
and can be counted. The genu and tibia have about six to fifteen
dorsal nude setae. The tarsi of legs II, III, and IV have fewer
nude setae and occasionally lack nude setae altogether.

Audy (1954:128) distinguished several types of nude setae, from
blunt, striated, nearly flask-shaped setae to attenunated, unstriated
setae, and noted the occurrence of microsetae. These types ( except
microsetae) tend to grade into one another in the material ex-
amined by the writer. Nude setae of the more proximal segments
tend to be unstriated and attenuated, although blunt striated setae
are not excluded. Tarsi II, III, and IV usually have at least one

182 The Uni\'ersity Science Bulletin

long untapered striated seta reminiscent of the tarsala of the larval
stage; however, the seta may be absent and often more than one is
present. Another feature of these tarsi is a slight recurved seta
on the dorsal distal margin, which resembles the subterminala of
the larval tarsi I. These setae are not at all constant in occurrence.
Variation in nude setae appears to be of such magnitude as to re-
quire a precise tabulation such as that proposed by Audy, if charac-
ters involving these setae are to be found.

Useable characters no doubt exist in leg setation but the most
likely ones appear to be those involving counts of setae on the more
proximal segments, rather than features of the types of nude setae.
At any rate a considerable amount of laborious tabulation is needed
before these features can be utilized.

RELATIONS OF MORPHOLOGY TO HABITS AND

ENVIRONMENT

Little is known of the habits and natural habitats of postlarval
trombiculids, since the postlarval stages are not often collected.
Loomis (1956), however, gives what he believes to be the habitat
of the postlarval stages for most of the species considered in the
present paper.

Trombiculid postlarvae are dwellers in soil or in similar sub-
strates. The habitats suggested by Loomis include grassland and
woodland soils, mammal nests or burrows (and soil associated
with these), decaying wood, crevices in rocky outcrops, and bat
caves. Quite possibly all of these habitats provide a substrate
of similar physical conditions (perhaps a somewhat granular sub-
strate in which the mites can burrow) so that little adaptation to
the physical makeup of the substrate of different habitats is needed.
At any rate there are no consistent morphological differences be-
tween species in different habitats. Species which occur in de-
caying logs (Trombictila splendens, for example) may be morpho-
logically similar to close relatives which occur in soils (in this case
Trombictila alfreddugesi) . Larvae of Trombicula merrihewi have
been taken only from bats, and larvae of T. ornata have been found
only on rodents, yet the nymphs of these two species are quite
similar. From this it appears that the postlarval habitat as sug-
gested by larval host preferences, does not provide sufficient in-
formation to permit conclusions concerning the relationships of
morphology to habitat.

Nymphs of the Trombiculidae 183

Additional information is available from seasonal occurrence and
observations on cultures. Since some species show marked dif-
ferences in seasonal abundance in the larval stage, it may be as-
sumed that their postlarvae have different temperature require-
ments and tolerances. Within the genera Trombictda and Eu-
schongastia, nymphs of winter species of chiggers tend to have
stouter cheliceral bases and wider cheliceral blades than do nymphs
of summer chiggers. There are some exceptions to this tendency.
It seems unlikely that this modification is directly associated with
temperature tolerances.

Observations on cultures indicate that nymphs of similar species
have different humidity requirements (Wharton and Fuller, 1952:
149). If there is a relationship between morphological features
and humidity requirements in trombiculid nymphs, it is not an ob-
vious one. The postlarvae of Whortonia senase seemingly occur
in very dry habitats, and their leaflike body setae possibly provide
protection against dessication. Such modifications are not seen
among other nymphs which may inhabit very dry situations.
Species of Hannemanio and of Blankoartia, whose postlarvae must
exist in very moist habitats, show no features which are clearly
adaptations to such environments.

Gross culture observations disclosed no differences in behavior
between postlarvae which have eyes and those which lack eyes.
Nevertheless, it happens that those postlarvae which are not un-
commonly collected in the field are those which have lenses (Blan-
kaartia in Panama and Etitrombicula in many regions); possibly
these species more readily come to the surface and are detected
by collectors. This does not hold true for species in the subfamily
Leeuwenhoekiinae, nymphs of which usually have lenses but are
not commonly collected.

The food material taken by numphs in culture offers additional
information concerning their habits. Although data are not avail-
able for all species, the general picture is as follows. Insect (Col-
lembola) eggs only: Euschongostia, Tromhicida (and probably
Blankoartia and Speleocola), and Hannemania. Collembolans only:
ChcJailonta, Neoscliongastia, P.seudoschongastia, and Walchia. Did
not feed: Eiischongastoidcs. Information is inconclusive for the
remainder of the genera (all leeuwenhoekiines ) . Some specimens
of Trombictda occasionally eat active collembolans.

Those species which eat active prey show gnathosomal modi-
fications possibly associated with their food habits. The genera

184 The University Science Bulletin

Chehidonta and Neoschongastia, particularly the former, have
elongated gnathosomal elements which possibly assist in the cap-
ture of prey. The genera Psetidoschongastia and Walchia, how-
ever, have shortened gnathosomal elements. These latter genera
have a more heavily sclerotized pharynx with reticulate markings,
which suggests more powerful musculature.

COMPARISON OF LARVAL AND NYMPHAL MORPHOLOGY

This section includes an abbreviated account of larval morphol-
ogy for comparison with the nymphal morphology. Emphasis is
placed upon those forms for which the nymphal morphology is
described. More detailed accounts of larval morphology may be
found in Wharton and Fuller (1952:30) and, for Kansas materials,
in Loomis (1956:1223).

The larva is six-legged rather than eight-legged but otherwise
is essentially similar to the nymph in gross morphology. The body
is at most only slightly constricted; it is capable of distention dur-
ing engorgement. There is no closed sternum between legs I.
The genital opening is absent. The scutum is not modified into a
linear stricture but retains its platelike nature. Rarely caudal
plates are present on the posterior dorsum of the abdomen.

The cheliceral base is short and the blade pointed. In most
species teeth of the cheliceral blade are restricted to the "tricuspid
cap," which consist of a dorsal tooth, a ventral tooth, and the point
of the blade. In species of Hannernanio the cheliceral blades are
expanded distally and provided with additional teeth. Species
of Chelandonta have a ventral row of minute serrations on the
cheliceral blade. Only species of Acomatacarus and Whartonia
have teeth on the dorsal margin of the blade, as do most nymphs.

Palpi of larvae are of rather similar shape. There is no distinct
trochanter. The setal arrangement is mostly constant. There
is one dorsal seta on the femur, and one dorsal seta on the genu;
the tibia has one dorsal, one lateral, and one ventral seta. The
condition of these palpal setae (whether nude or with branches)
is a specific character. There are no accessory claws; the major
claw is present and is presumably always branched, thus differing
from that of the nymph. The number and arrangement of these
branches is a specific character and to some extent a group charac-
ter. The vestiture of the palpal tarsus is variable but has been
little studied.

Nymphs of the Trombiculidae 185

Other features of the gnathosoma have been Httle used. The
galeal setae are seemingly homologous to the apical hypostomal
setae of the nymph. The galeae are paired processes which appear
to curve dorsally around the chelicerae. Each bears a seta; the
characteristics of these setae are used as specific characters.

The scutal plate contains the characters most used in larval clas-
sification. Basically this is a rectangular plate bearing several
setae. One anteromedian seta is present in most species (paired
in Leeuwenhoekiinae, absent in Walchia). The four comers each
bear a seta in most species, but in Pseudoschongastia the postero-
lateral setae are off the scutum and Walchia bears additional pos-
terior setae. Scutal shapes are correspondingly modified; the
scutum is shortened in Pseudoschongastia and lengthened in Wal-
chia. The shape of the scutum has been used as a specific character
and sometimes as a group character. Most species of Etischongasiia
have broad short scuta. Species of Trombictda have scuta which
vary from rectangular to pentagonal, with the apex directed posteri-
orly. In Neoschongastia the scutum is partially submerged beneath
the cuticular striae. Leeuwenhoekiine genera (except Whartonia)
have a small anteromedian projection on the anterior margin. The
sensilla vary from flagelliform to expanded. The nature and degree
of branching is used as a specific character among those species
with flagelliform sensilla. When expanded, the degree of expansion
is used as a specific character; these sensilla are all more markedly
enlarged than any of the sensilla of the postlarval stages. Ex-
panded sensilla are characteristic of the genera Cheladonta, Eu-
schongastia, Euschdngastoides (one of two species), Neoschongastia,
and Pseudoschongastia. Scutal markings associated with the sen-
sillary bases are used as specific characters. A series of ten "stand-
ard" measurements taken on the scutum are used as specific charac-
ters. Scutal modifications in the larval stages do not seem to be
related to modifications in the nymphal stage.

Shapes and details of branching of body setae have been used
as specific and sometimes generic characters; usually these setae
are not so strongly modified as are those of the nymphs, and the
characters have not been extensively used. The body setae of the
larvae are arranged in rows and can be easily counted; their num-
ber and arrangement are specific characters.

Some group and specific characters are taken from the legs. Legs
contain six or seven segments, depending on the division of or fusion

186 The University Science Bulletin

of the femur. In the leeuwenhoekiines all legs have six segments.
Trombiculine species have seven segments in leg I and usually
seven in legs II and III; Walchia and Psciidoschdngastia clearly
have six segments in legs II and III, and it is difficult to determine
whether the femoral division is complete in some other species.
The setation of the legs is sparse and constant, so the number and
nature of leg setae is a widely used specific character.

Characters which retain similar states in both larval and post-
larval stages would be of considerable interest, since it would be
possible to predict the characters of one stage from those of the
other. However, such characters are very few. Sasa (1953:429)
mentions three features which are similar in larva and postlarva,
these being color, nature of the basal part of the sensillum, and
number of anteromedian and tectal setae. The present writer must
find fault with the first two of these.

The color shown by trombiculids is undoubtedly influenced by
environment and by age, as may be seen in cultures. Although
some species show little color variation (for example, species of
Pseudoschongostia and Walchia) others show a wide range. Lar-
vae of Tromhicida montanensis range in color from white to dark
orange; nymphs range from yellow to orange. The writer has
placed larvae in culture which he characterized as "yellow"; nymphs
produced from these larvae were characterized as "orange." On
the other hand, Loomis (personal communication) has collected
and cultured adults of Tromhicida splendens (normally red) which
were white; larvae obtained from these adults were also white.
Thus, while the color of larval and postlarval stages is related and
may be similar, variation is great enough that the writer would
hesitate to predict the color of one stage from that of the other.

Since a character that has been emphasized in studies of the
larval stage is the sensilla (whether flagelliform or expanded) any
similar or related modifications of the sensilla of the postlarval
stages would be of greatest interest. Unfortunately no such related
modifications were found; it is not possible to tell from the sensilla
of the nymph whether the larva had flagelliform or expanded sen-
silla. Sasa observed that some species of Lcptotromhidium have
similar short basal branches on the sensilla in both larval and post-
larval stages. This must be regarded as an exceptional case; no
constant relationship appears among the species studied by the
v^iter.

The number of anteromedian setae on the larval scutum is di-
rectly related to the number of tectal setae on the adult scutum, as

Nymphs of the Trombiculidae 187

reported by Sasa and by others. Leeuwenhoekiine species have two
tectal setae and two anteromedian setae; species of Wolchia have
no tectal setae and no anteromedian setae, while other trombiculines
have one tectal seta and one anteromedian seta. The nature of the
tectal setae (length, modification of stem, and details of branching)
are perhaps related in some species but not in most.

Other features occasionally show similarities in larva and post-
larva for some species. For example species with large larvae often
have large nymphs, but this relationship is far from constant.

Trombiculid mites are certainly not unique in this exhibition
of independent variation of characters in two differently adapted
stages. Where two stages lead different modes of existence, such
independent variation is an obvious advantage, since characters
selected for in one stage need not be exposed to adverse selection
in subsequent stages. Seemingly natural selection has built up a
group of genes which do not express themselves (or have little
expression) in the larval stage but which have their major eflFects
in the postlarval stages. The metamorphosis of holometabolous
insects is a similar and better-known phenomenon.

When considering these problems it is necessary to bear in mind
constantly that the larva, nymph, and adult of an individual are
all the same animal; that there is but one genotype for the indi-
vidual; and that the effects of selection on the characters of any
one stage will be felt upon the entire genotype.

TAXONOMY OF THE NYMPHAL STAGE

Nearly all of the species and group names used in this study are
based upon the larval stage. Only about half of the established
genera are known as postlarvae, and but a small fraction of the
described species are known as postlarvae. These circumstances
make difficult the construction of group diagnoses and keys; one
is hesitant to characterize a genus on a few species when he is
aware of the existence of many more species, and perhaps he has
not even seen the type of the genus, wliich he certainly should con-
sider. Womersley ( 1952 ) has provided keys to genera and species
but usually omits a formal diagnosis for groups, Sasa's (1952) ex-
cellent work lacks even keys, and Wharton and. Fuller (1952) in-
clude only the postlarval characterizations of some groups. The
present writer has chosen to use the species at hand as the basis
for a full description for each genus, including most characters
common to all included species. These descriptions will undoubt-
edly require modification as more postlarvae are described, but

188 The University Science Bulletin

they may provisionally serve the purpose of a source for critical
comparison. In the construction of these descriptions the previously
described species were considered; however, as most forms are
inadequately known, it was not possible to tell how accurately these
fit the generic descriptions. In certain cases previously described
species do not agree with the generic characters. These species are
provisionally placed in genera but the generic diagnoses vdll ob-
viously have to be modified when these species are better known
and can be classified with greater certainty.

The keys include most of those genera known as postlarvae, but
some are inadequately described and were not placed in the keys.
These keys were designed primarily for nymphs but will probably
classify most adults as well. Keys to species are given; however,
in most cases it was possible only to include those species seen by
the writer. These specific keys will have little value except in the
midwestern region of the United States, but it is hoped that they
can be of use also in the study of other postlarvae when they are
known.

In the generic diagnoses, characters are numbered and are pre-
sented in the same sequence for each genus; the numbers facilitate
comparisons and discussions. Within each subfamily each char-
acter retains its number for all generic descriptions, but the systems
are different for the two subfamilies. Particularly important char-
acters in generic recognition are italicized.

In descriptions, the synonomy of each species is not complete, but
includes only the original citation, some different name combina-
tions, and important references to larva or postlarva. Complete
synonymies for most included species may be found in Wharton
and Fuller (1952) augmented by Loomis (1956).

Group characters are not repeated in the specific descriptions.
Measurements are given in microns, except where indicated; means
followed by their standard errors are used where possible. Setal
counts on the palpal tibia and tarsus include only branched setae;
the nude seta at the base of the palpal claw, the tarsala, and the
apical nude setae of the tarsus are not included in these counts.

In the listing of specimens examined, a uniform procedure was
followed similar to that of Loomis ( 1956 ) . First is given the total
number of nymphs examined by the writer. Then follows the col-
lection data pertaining to the larval hosts of these nymphs, in the
following order: State, county, exact locality (where known), host,
and the date host was taken. The collector of the host is omitted
in most cases; usually these animals were taken by the University

Nymphs of the Trombiculidae 189

of Kansas Chigger Project and were either obtained by Richard B.
Loomis or catalogued by him. Lastly, for each collection the num-
ber of nymphs examined is given, with an indication of the persons
who reared them. Workers on the Project are identified by initials,
as follows: DAC: D. A. Crossley, Jr.; EHK: Ervin H. Kardos, LJL:
Louis J. Lipovsky; RBL: Richard B. Loomis. Also, where nymphs
were identified by recovering their larval skins, this is indicated in
parentheses. Otherwise, nymphs were identified by association of
larvae or by comparison with other nymphs.

Family Trombiculidae Ewing
Ewing, 1944, Proc. Biol. Soc. Washington, vol. 57, p. 101.

Diagnosis (nymph). — Body small to medium in size (about
0.4-1.2 mm.); white to red in color; constricted or not; oval to some-
what elongated; provided with numerous branched or modified
setae. Eyes sometimes present, 1/1 or 2/2, anterior to or adjacent
to sensillary area. A rectangular to pentagonal closed sternum
usually present between anterior coxae; sternum sometimes open
behind, precoxal plates of coxae I sometimes present; sternal area
sclerotized or not. Genital opening bet\veen coxae IV; genital
plates bearing six genital setae in addition to body setae; only two
pairs of genital suckers. Anus posterior to genital opening, divided
or not. Legs in two groups; each leg of seven segments and bearing
a pair of claws; both branched and nude setae numerous on legs,
particularly legs I; tarsus I with or without apical stumplike process.

Gnathosoma with basis capituli evident as a sclerotized ring,
bearing hypostome, palpi, and chelicerae. Pharynx usually indis-
tinct but rarely visible due to reticulate markings. Hypostome
usually pointed, projecting; occasionally blunt; bearing about eight
usually nude, apical setae or about twenty nude or nearly nude
apical setae. Chelicerae composed of base, blade, and incon-
spicuous pseudochela; base stout to elongated; blade usually shear-
like, sometimes daggerlike or strongly curved, usually with dorsal
teeth or serrations. Palpi composed of five segments: trochanter
small, without setae; femur large, with branched setae; genu large
but smaller than femur, with branched setae; tibia small, somewhat
pointed, bearing claw at apex and usually two subapical accessory
claws, several branched setae, and a nude seta on lateral surface at
base of claw; tarsus small, articulated to ventral surface of tibia,
bearing several usually branched setae, one tarsala, and one to
several short apical nude setae.

190 The University Science Bulletin

Scutum composed of anterior membranous tectum, rodlike crista,
and posterior sensillary area. Tectum large to small; usually
hyaline but sometimes slightly sclerotized; margin smooth or bear-
ing teeth or serrations; with one, two, or no tectal setae. Crista
usually a thin rod, sometimes very thin or broadened; sometimes
with prominent punctation. Sensillary area variable in shape, from
circular to transverse-oval or triangular; variously modified; bearing
two sensilla, these flagelliform to subclavate and usually with
branches; area usually surrounded by ridges; usually with a pos-
terior apodeme.

Remarks. — The combination of bladelike chelicerae (never
elongate-stylettiform ) , no more than two tectal setae, and a single
pair of sensilla on the scutum, will separate trombiculid nymphs
and adults from most of the genera in the trombidioid complex.
The constricted, "figure 8-shaped" body is not characteristic of all
genera but is nevertheless a useful field character.

Extensive larval diagnoses have been given by Wharton and
Fuller (1952:40) and by Loomis (1956:1229).

Family rank is accorded these mites by most workers, although
some prefer a subfamily rank in the family Trombidiidae.
Womersley recognizes two families, Trombiculidae and Leeuwen-
hoekiidae. The proper position must, of course, be decided by
comparison of the chiggers with their closest relatives, which are
obviously the trombidiids. Such a comparison has been quite diffi-
cult, since trombidiids are mainly known from their adult stage,
while trombiculids are mainly known as the larval chiggers. More
recent studies of reared material have increased knowledge of
both groups and the result is an increasing difficulty in separating
them. The trombidiid genus Neotromhidium, for example, is
morphologically more similar to leeuwenhoekiine genera than to
most trombidiid genera. The biological character of vertebrate
parasitism by trombiculids and invertebrate parasitism by trom-
bidiids will exclude these questionable genera from Trombiculidae;
they may be excluded on morphological grounds by a rigid and
perhaps artificial definition such as that given above. However,
these techniques do not solve the problems of the limits of the
trombiculid group or place them in their proper taxonomic position.

At the present time the best solution appears to be retention
of the family rank for trombiculids, until such time as the ac-
cumulated evidence either verifies the position or forces a change
to subfamily (or superfamily) status. Knowledge of trombidiids
is still fragmentary; trombiculids are better known but still require

Nymphs of the Trombicijlidae 191

extensive taxonomic investigation at the higher levels. The evidence
for change at the present must be judged inconclusive.

Suhfamilies. — Most workers divide the Trombiculidae into four
subfamilies, these being Trombiculinae, Walchiinae ( = Gahrlie-
piinae), Leeuwenhoekiinae, and Apoloniinae. The two latter sub-
families Womersley places in Leeuwenhoekiidae. As no species of
Apoloniinae are known as postlarvae, this subfamily must be
omitted from the following discussions.

The subfamily Walchiinae has recently become a trouble-spot
in chigger taxonomy. Wharton and Fuller (1952:41) separate
Walchiinae from Trombiculinae in the larval stage by the number
of segments in legs II and III; Walchiinae have but six segments,
the femurs being undivided, while trombicuHnes have seven seg-
ments due to the division of the femora into basifemur and telo-
femur. Some workers, including the writer, have experienced dif-
ficulty in using this character. Womersley (1952:278) resolved the
problem by removing to Trombiculinae the genera Pseudoschon-
gastia and Walchiella, and considering the remaining genera con-
generic with GalirUepia (with the subfamily name Gahrliepiinae).
Larvae were diagnosed by the absence of the anteromedian scutal
seta (present in Trombiculinae) and postlarvae were recognized
by the presence of an apical stumplike process on tarsus I (absent
in Trombicuhnae). Audy (1954:161) and Loomis (1956:1362)
followed Womersley's arrangement but expressed doubts as to the
propriety of recognizing only the genus GalirUepia.

The writer's studies of postlarval stages show that Walchia is
closely similar to bo!h Pseitdoschdngastia and Eiischongastoides; it
appears that a subfamily would have to contain all three of these
genera. However, such a subfamily would be difficult to diagnose
on the characters based on the larval stage, since the included
species would combine such features as anteromedian scutal seta
present or absent, scutum reduced (Pseudoschdngastia) or extended
(Walchia), sensilla globose or flagelhform (E. hoplai), and legs II
and III six- or seven-segmented. Also, larvae of species of
Euschongastoides and perhaps Pseudoschdngastia are very similar
to some imdoubted trombiculines. Nevertheless, the impression
remains that these three genera form a natural group. It may be
that further studies on the larval stage will produce characters
which justify the recognition of a subfamily in spite of the diversity
in that stage. Until this is accomplished the subfamily Walchiinae
(or Gahrliepiinae) must be abandoned as a synonym of
Trombiculinae.

192 The University Science Bulletin

Key to Subfamilies of Trombiculidae (Nymphs)

1. Tectum with two setae, usually side-by-side; sensilla long, flagelli-
form, without branches; setae on palpal genu less numerous than
those on palpal femur Leeuwenhoekiinae p. 192

1'. Tectum with one seta or without setae; sensilla variable, with at
least short basal branches ( except Blankaartia ) ; setae on palpal
genu more numerous than those on palpal femur, or the same num-
ber on those segments Trombiculiiiae p. 205

Subfamily Leeuwenhoekiinae Womersley
Womersley, 1944, Trans. Roy. Soc. South Australia, vol. 68, p. 102.

Diagnosis {nijmj)Jis). — Body oval, not constricted (except Hanne-
mania). Tectum with two setae, usually side-by-side. Sensilla long,
flagelliform, without branches. Anus circular, not obviously di-
vided into two shoelike portions. Setae on palpal genu less nu-
merous than those on palpal femur.

Additional features (nymphs). — Body with eyes present or ab-
sent, 1/1 or 2/2, anterior to and separated from sensillary area;
sternal area sclerotized but no true sternum formed by a crossbar
connecting coxae II; precoxal plates never present. Gnathosoma
with cheliceral base of usual shape (mean ratio BL/BH: 1.8-2.4),
cheliceral articulation with basis capituli ventral; two accessory
claws on palpal tibia; pharynx inconspicuous, not marked with re-
ticulate pattern. Scutum with tectal margin smooth; mean ratio
ASL/SB: 3.2-5.1; bulla absent; lateral keels absent; carina absent;
basal plate of two stiaplike bands; only posterior and lateral ridges
on sensillary area; apodeme present.

Included genera. — Known from both larvae and postlarvae:
Acomatacarus Ewing, Chatia Brennan, Hannemania Oudemans,
Leeuwenhoekia Oudemans, and Whartonia Ewing. Knowoi from
larvae only: Odontacarus Ewing and Shunsennia Jameson and
Toshioka.

Reiruirks. — The included genera are substantially separable from
the trombiculine genera, so that the subfamily status is merited.
As previously mentioned, Womersley (1945:47) has given the
group full family rank, but was not followed by Wharton and
Fuller (1952:96). The present writer also prefers to retain the sub-
family rank.

The genus Odontacarus (known only as larva) contains but two
species. According to Wharton and Fuller (1952:103), "the
specimens in existence are too badly damaged to study satis-
factorily." "Probably a synonym of Acomatacarus (Acomatacarus) ."

Nymphs of the Trombiculidae 193

Nymphs of Shunsennia have been reared and are being described
by Lipovsky (personal communication). All other genera are
known as postlarvae, but three of five subgenera of Acomatacarus
are known as larvae only.

Leeuwenhoekiine larvae are found on amphibians, reptiles, birds,
and mammals. The postlarvae are apparently adapted to various
habitats.

Key to Geneka of Leeuwenhoekunae (Nymphs)

1. Body constricted; posterior body setae elongated, each ending in a
hooklike spine Hannemania p. 200

r. Body not constricted; posterior body setae not ending in hookUke
spines 2

2. Posterior body setae with simple branches; setal tips single, long,
attenuated Chatia p. 198

2'. Posterior body setae modified, not ending in single, long attenu-
ated tips 3

3. Posterior body setae leaflike Whartonia p. 203

3'. Posterior body setae somewhat expanded 4

4. Tarsus longer (mean ratio TL/TH: 6.2), cylindrical in shape

Leeuwenhoekia p. 193
4'. Tarsus shorter (mean ratio TL/TH: 2.1), rather pyriform in shape

Acomatacarus p. 195

Genus Leeuwenhoekia Oudemans

Leeuwenhoekia Oudemans, 1911, Ent. Berichten, vol. 3, p. 137 (type Hetero-
thromhhim verduni Oudemans).

Diagnosis (nymph). — (1) Body shape, oval, not constricted. (2)
Posterior body setae modified, expanded. (3). Ratio TL/TH about
6.2. ( 4 ) Tarsus I cylindrical.

Additional features (nymph). — (5) Body size medium (about
0.8 mm. long). (6) Eyes 1/1. (7) Adjacent body setae of similar
lengths. (8) Sclerotized sternal plate present. (9) Legs not unusu-
ally elon^^ated; branched setae slightly expanded. (10) Cheliceral
blade shearlike. (11) Palpus moderate in size; claw moderate in
size; tarsus with about nine branched setae and six or seven apical
nude setae. ( 12 ) Hypostome pointed, projecting, with about eight
apical nude setae and about twenty ventral branched setae. (13)
Basis capituli of normal shape. (14) Tectum large, well sclerotized,
saggitate in .shape; tectal setae elongated, not expanded. (15)
Crista a narrow rod. ( 16) Sensillary area oval; mean ratio ASL/SB:
5.1. (17) About ten to fifteen parascutal setae.

Remarks. — Wharton and Fuller (1952:96) recognize but two
species of Leeuwenhoekia in two subgenera; these are verduni

7—8920

194 The University Science Bulletin

(Oudemans) in the typical subgenus and americana (Ewing) in the
subgenus comatacarus Ewing. Since verduni has not been reared,
the above description is based upon americana only.

Leeuwenhoekia as represented by americana is very similar to
Acomatacarus (Acomatacartis) ; these two may be separated by the
shape of tarsus I as evidenced by characters ( 3 ) and ( 4 ) but these
are of doubtful generic significance (some postlarvae of Acomata-
carus, as described by Womersley, probably key to Leeuwenhoekia ) .
If americana is really congeneric with verduni, the Acomatacarus is
probably a synonym of Leeuwenhoekia.

Leeuwenhoekia americana (Ewing)

(Figs. 18, 62, 96, 97, 140, 185, 197)

Comatacarus americanus Ewing, 1942, Jour. Parasit., vol. 28, p. 490 (larva).
Leeuwenhoekia (Comatacarus) americanus, Wharton and Fuller, 1952, Mem.

Ent. Soc. Washington, no. 4, p. 96 (larva); Loomis, 1956, Univ. Kansas Sci.

Bull., vol. 37, pp. 1234-1235 (larva).

Description of nymph. — Body: About 0.8 mm. long, color yellow.
Eyes red in life, about 12 \i in diameter. Setae expanded; tips
rounded; branches thick; posterior setae about 20 \i long. Sternal
area containing nearly square sclerotized plate bearing 8 setae.
Measurements of tarsus I (1 specimen); TL: 185, TH: 30, TL/TH:
6.2.

Gnathosoma: Cheliceral base stout, with conspicuous puncta;
blade with about 15 flat teeth. Cheliceral measurements (1 speci-
men); BL: 109, BH: 58, CL: 66, BL/BH: 1.9, BL/CL: 1.7. Pal-
pus with outer femoral and outer genual setae bearing short paired
branches rather than longer single branches; tarsus with 4 inner, 2
outer apical nude setae; tarsala about 13 jj, long. Setal counts (1
specimen); femur: 15.5, genu: 10, tibia: 6, tarsus: 9.

Scutum: Tectal setae apical. Crista with several puncta. Basal
plate broad; apodeme narrow and elongated; sensillary bases
towards posterior of sensillary area. Scutal measurements ( 1 speci-
men); ASL: 107, SB: 21, TS: 43, SENS: about 60, ASL/SB: 5.1.

Remarks. — On the single occasion on which this species was
reared, one of two larvae reached the nymphal stage. ( The nymph
was offered collembola eggs but did not eat.

Specimens examined. — One nymph, as follows: Kansas. Chey-
enne County: 15 mi. N, IIM W St. Francis, Peromijscus maniculatus,
November 1, 1952 ( 1 nymph, reared DAC ) .

Nymphs of the Trombiculidae 195

Genus Acomatacanis Ewing

Acomatacarus Ewing, 1942, Jour. Parasit., vol. 28, p. 490 (type Acomatacarus
arizonensis Ewing).

Diagnosis (nymph). — (1) Body shape oval, not constricted. (2)
Posterior body setae modijied, expanded. (3) Mean ratio TL/TH:
about 2.1. (4) Tarsus I rather pyriform.

Additional features (ntimph). — (5) Body size small to medium
(about 0.4-0.6 mm. long). (6) Eyes absent or 1/1. (7) With or
without long and short body setae intermixed. (8) Sclerotized
sternal plate present. (9) Legs not unusually elongated; branched
setae slightly expanded. (10) Cheliceral blade shearlike or dagger-
like. (11) Palpus short, rather small; some femoral setae expanded,
some genual setae expanded; claw moderate in size; tarsus with
about six to eight branched setae and three to five apical nude setae.
( 12 ) Hypostome pointed and projecting, or blunt; six to eight apical
setae, nude or branched; about ten ventral branched setae. (13)
Basis capituli normal to broad and short in shape. (14) Tectum
well sclerotized, intermediate in size; tectal setae expanded or not.
(15) Crista rodlike, narrow or somewhat broadened. (16) Sensil-
lary area oval to transverse-oval; mean ratio ASL/SB: 3.2-4.5". (17)
About ten to fifteen parascutal setae.

Remarks. — Womersley (1945:98) described the reared nymphs
of Acomatacarus ausfralensis (Hirst), A. longipes (Womersley),
and A. nova-quinea (Womersley). He also placed in that genus
Rhyncholophus retentus Banks, Dromeothrombium dromus Wom-
ersley, and Acomatacarus patrius Womersley, all known only as
adults. Their affinities with Acomatacarus were established by com-
parisons with the reared nymphs mentioned above. This action ap-
pears to have been well taken. The diversity of forms suggests that
eventually some generic reassignments will prove necessary but
for the present a broad genus Acomatacarus seems advisable.

The descriptions of these species as given by Womersley are gen-
erally in agreement with the generic description proposed by the
writer. One species, dromus, has long unmodified body setae.
Womersley reports that all of the species have no accessory palpal
claws but instead three or four short spines at the base of the main
claw. This observation should be verified.

In addition to the above mentioned species and the two de-
scribed in this paper, some 35 species of Acomatacarus known only
from the larval stage have been described.

196 The University Science Bulletin

Key to Subgenera of Acomatacarus (Nymphs)

1. Cheliccral blade shearlike A. (Acomatacarus) p. 196

1'. Cheliceral blade narrow, daggerlike A. (Xetuicarus) p. 197

Subgenus Acomatacarus Ewing

Acomatacarus, Womersley, 1945, Trans. Roy. Soc. South Australia, vol. 69,
p. 98 {nymph); Wharton and Fuller, 1952, Mem. Ent. Soc. Washington,
no. 4, p. 97 (nymph).

Diagnosis (7iymph). — (10) Chelicera with blade shearlike. (11)
Palpal tarsus with five apical nude setae. (12) Ihjpostome pointed,
projecting, with about eight apical nude setae. ( 14 ) Tectal setae
not expanded.

Remarks. — These features appear to be the ones of subgeneric
importance which characterize A. (Acomatacarus) as opposed to
A. (Xenacarus). The postlarvae described by Womersley appear
to fall within the typical subgenus.

Acomatacarus arizonensis Ewing

(Figs. 37, 142, 209)

Acomatacarus arizonensis Ewing, 1942, Jour. Parasit., vol. 28, pp. 490-491
(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1236-1238
( larva ) .

Description of nymph. — Body: about 0.4 mm. long, color dull red.
Eyes absent. Setae expanded; tips rounded; branches thin and
short; most setae short but longer ones (2-3 times longer) scat-
tered among these; posterior setae (longer ones) about 30 \x. long.
Sternal area with broad plate bearing ten modified setae. Measure-
ments of tarsus I (1 specimen); TH: 36, TL: 75, TL/TH: 2.1.

Gnathosoma: Cheliceral base stout, rather rounded, punctate;
blade with about 16 small teeth; apodeme of blade not constricted,
and blunt. Cheliceral measurements (1 specimen); BL: 56, BH: 31,
CL: 36, BL/BH: 1.8, BL/CL: 1.6. Palpus somewhat stout; outer
femoral and outer genual setae slightly expanded; tarsus with 3
inner, 2 outer apical nude setae; tarsala not seen. Setal counts ( 1
specimen); femur: 9-10, genu: 5-6, tibia: 5, tarsus: 8.

Scutum: Tectal setae apical. Crista rather broad. Sensillary
area with apodeme broad; sensillary bases small; basal plate wide.
Scutal measurements (1 specimen); ASL: 58, SB: 18, TS: 20, SENS:
about 60, ASL/SB: 3.2.

Remarks. — Several additional attempts to rear this species were
unsuccessful, possibly due to inadequate moisture control. The
species appears to be restricted to arid situations (see Loomis,
1956:1237).

Nymphs of the Trombiculidae 197

A. arizonensis lacks striking distinctive features; most of the
known postlarvae of Acomatacarus are clearly congeneric with
arizonensis ( the genotype ) , on the basis of their descriptions.

In the key given by Womersley (1945:110), arizonensis (as
nymph) will key out to dromus (known only as adult). These
species may be separated by the form of the body setae; in ari-
zonensis the dorsal body setae are expanded while in dromus they
are slender.

Specimens examined. — One nymph, as follows : Arizona. Cochise
County: Sceloporus jarrovii, July 8, 1951, taken by H. S. Fitch
(1 nymph, reared LJL).

Subgenus Xenacarus Greenberg

Xenacarus Greenberg, 1951, Jour. Parasit., vol. 37, p. 525 (type Acomatacarus
(Xenacarus) plumosus Greenberg).

Diagnosis (nymph). — (10) Chelicera with blade straight, dag-
gerlike. (11) Palpal tarsus with three apical nude setae. (12)
Htjpostome blunt, with about six to eight apical, sparsely-branched
setae. (14) Tectal setae somewhat expanded.

Remarks. — Of these features, the blunt hypostome and the
straight, daggerlike cheliceral blade appear to be characters of
group importance. The expanded tectal setae are of interest.
Other species with expanded or modified body setae nevertheless
have slender tectal setae.

Acomatacarus plumosus Greenberg

(Figs. 34, 88, 134, 135, 138, 189, 205, 207)

Acomatacarus plumosus Greenberg, 1951, Jour. Parasit., vol. 37, pp. 525-527
(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1240-1241
( larva ) .

Description of nymph. — Body: About 0.6 mm. long, color white.
Eyes 11; red in life; about 12 \). in diameter. Setae with body ex-
panded but base of normal size; tips rounded; branches of two types,
mostly large and heavy but with small branches interspersed; adja-
cent setae of similar lengths; posterior setae about 25 ix long.
Sternal area with very broad plate bearing about ten modified
setae. Measurements of tarsus I (means of 9 specimens); TL:
124.7 ± 2.789; TH: 58.2 ± 1.025, TL/TH: 2.13 ± 0.0289.

Gnathosoma: Chelicera with base rather stout, punctuate; blade
with about 12 small flat teeth; apodeme of blade not constricted.
Measurements of cheliccrae (means of 9 specimens); BL: 60.7 db
0.928, BH: 32.9 ± 0.605, CL: 37.9 ± 0.754, BL/BH: 1.84 ± 0.041,

198 The University Science Bulletin

BL/CL: 1.61 ± 0.0602. Palpus somewhat stout; outer femoral
and outer genual setae expanded; tarsus with 2 inner, one outer
apical nude setae; tarsala 8 \). long. Setal counts; femur: 10.56 ±
0.448 (8 specimens), genu: 6.8 ± 0.214 (10 specimens), tibia:
range 4-5, tarsus: range 6-7.

Scutum: Tectal setae apical; expanded, resembling body setae.
Crista narrow. Sensillary area oval; basal plate narrow; apodeme
narrow. Scutal measurements (means of 8 specimens); ASL: 104.8
± 3.98, SB: 23.7 ± 0.799, TS: 26.3 ± 2.496, SENS: about 100,
ASL/SB: 4.53 ± 0.1878.

Remarks. — This species was reared to the nymphal stage on sev-
eral occasions. Collembola eggs were offered as a food material
and the nymphs possibly ate the eggs or the freshly-hatched col-
lembolans; feeding was not observed. It is not certain that they fed
at all.

A. phimostis is readily identified by the form of the body setae.
In other respects it also differs markedly from arizonensis, and is
probably generically distinct. The one other species of Acomata-
carus (Xenacanis), A. brevicalar Brennan and Jones, is known as
larva only.

Specimens examined. — Total 16 nymphs, as follows: Kansas.
Barber County: 4 mi. S Aetna, Neotoma micropus, August 22, 1949
(4 nymphs, reared LJL), September 15, 1953 (3 nymphs, reared
DAC, idet. by larval skin); 3/2 mi. S, 1 mi. W Aetna, Neotoma
micropus, April 11, 1949 (8 nymphs, reared LJL); 4 mi. S Aetna,
Peromyscus leucopus, September 15, 1953 ( 1 nymph, reared DAC ) .

Genus Chatia Brennan

Cliatia Brennan, 1946, Jour. Parasit., vol. 32, p. 132 (type Chatia setosa Bren-
nan); Wharton et al., 1951, Jour. Parasit., vol. 37. p. 30 {nymph); Audy,
1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 163 (nymph).

Diagnosis (nymph). — (1) Body shape oval, not constricted. (2)
Posterior body setae not modified, branched, and ending iti long,
single, attenuated tips; not expanded. (3) Ratio TL/TH: 2.5.(4)
Tarsus I cylindrical.

Additional features {nymph). — (5) Body size medium (about
0.8 mm. long). (6) Eyes 1/1. (7) Adjacent body setae of similar
lengths. (8) Sclerotized sternal plate present. (9) Legs unusually
elongated; branched setae not expanded. (10) Cheliceral blade
shearlike. (11) Palpus stout, large; claw somewhat elongated;
tarsus with about fourteen branched setae and six or seven apical
nude setae. ( 12 ) Hypostome pointed, projecting; with about eight

Nymphs of the Trombiculidae 199

apical nude setae and numerous ventral branched setae. ( 13 ) Basis
capituli of normal shape. ( 14 ) Tectum well-sclerotized, large, sag-
gitate in shape; setae elongated, not expanded. ( 15) Crista a broad
rod, with prominent punctation. (16) Sensillary area oval; ratio
ASL/SB: 5.1. (17) About ten to fifteen parascutal setae.

Remarks. — A few characters for nymphs of Chatia were given
by Wharton et al. (1951:30) and by Audy (1954:163). Most note-
worthy are the elongated legs (all longer than the body), the
prominent puncta of the crista, and the attenuated tips of the body
setae.

Chatia setosa Brennan
(Figs. 16, 92, 93, 141, 190, 210)

Chatia setosa Brennan, 1946, Jour. Parasit., vol. 32, p. 132 (larva); Audy, 1954,
Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 163 (nymph).

Description of nymph. — Body: About 0.8 mm. long, color not
known. Eyes about 23 [). in diameter. Body setae normal, with
rather short branches; tips long, single, attenuated; posterior setae
about 75 pi, long. Sternal area containing nearly square sclerotized
plate, bearing about 30 setae. Legs noticeably elongated, all longer
than body. Measurements of tarsus I (means of 3 specimens); TL:
266.3 ± 17.37, TH: 107.0 ± 2.517, TL/TH: 2.50 =h 0.2082.

Gnathosoma: Chelicera with base large, stout, punctate; blade
with about 35 to 50 small to tiny teeth, apodeme of blade not con-
stricted. Cheliceral measurements (means of 3 specimens); BL:
175.7 ± 1.33, BH: 75.0 ± 1.15, CL: 99.0 ± 1.00, BL/BH: 2.33
± 0.067, BL/CL: 1.80 ± 0.0000. Palpus with setae numerous;
tarsala about 13 \). long; tarsus with apical nude setae arranged 1
outer, 5 or 6 inner. Setal counts (means of 3 specimens); femur:
30.43 ± 1.00, genu: 20.00 ± 1.50, tibia: range 8-11, tarsus: range
13-18.

Scutum: Crista broadly joined to- sensillary area; apodeme large,
prominent; sensillary bases at back of sensillary area. Scutal meas-
urements (means of 3 specimens); ASL: 159.7 dr 5.84, SB: 32.5 ±
0.500, TS: 85.7 ± 4.63, SENS: about 120, ASL/SB: 5.10 ± 0.100.

Remarks. — This is the only species in the genus. The specimens
were reared by Dr. James M. Brennan, and were originally loaned
to Mr. Louis J. Lipovsky. Dr. Brennan and Mr. Lipovsky have very
kindly allowed me to describe them.

Specimens examined. — Total 3 nymphs, as follows: Montana.
Ravalli Cotmty: Citellus lateralis cinerascens, field no. AP-22355,
May 22, 1946 (2 nymphs, reared by Dr. Brennan); Tamiasciurus

200 The Unintersity Science Bulletin

hudsonicus richardsoni, field no. AP-21201, November 11, 1944 (1
nymph, reared by Dr. Brennan).

Genus Hannemania Oudemans

Hannemania Oudemans, 1911 Ent. Berichten, vol. 3, p. 137 (Type Hetero-
thromhium ht/lodeus Oudemans); Wharton et ah, 1931, Jour. Parasit., vol.
37, p. 30 (adult).

Diagnosis (nymph). — (1) Body constricted, figure 8-shaped.

(2) Posterior body setae elongated, not expanded, each ending in

hooklike spine. (3) Mean ratio TL/TH: 2.1-2.5. (4) Tarsus I

nearly cylindrical.

Additional features (nymph). — (5) Body size medium to large
(about 0.9-1.2 mm.). (6) Eyes 2/2. (7) Adjacent body setae of
similar lengths. (8) Sternal area sclerotized. (9) Legs not unusu-
ally elongated; branched setae not expanded. ( 10) Cheliceral blade
shearlike. (11) Palpus stout, size moderate to large; claw moderate
in size; tarsus with about ten to twenty branched setae and five
apical nude setae. (12) Hypostome pointed, projecting; with about
eight apical nude setae and numerous ventral branched setae. ( 13 )
Basis capituli normal to short and broad in shape. (14) Tectum
smxill, well-sclerotized; tectal setae elongated, not expanded. (15)
Crista a narrow rod. ( 16) Sensillary area oval; mean ratio ASL/SB:
3.7-4.5. ( 17 ) About fifty parascutal setae.

Remarks. — Larvae of species of Hannemania parasitize only am-
phibians; they are unique in that they burrow into the skin. The
larvae may also be recognized by their peculiar, expanded chelic-
erae.

Postlarvae are distinct from other leeuwenhoekiines particularly
in characters (1), (2), (6), (14), and (17) as indicated above. Of
these, the constricted body ( 1 ) is puzzling since this feature is typi-
cal of Trombiculinae. Otherwise, Hannemania is more closely sim-
ilar to other leeuwenhoekiines. Mounted specimens are readily
identified by the strong hooklike tips of the posterior body setae.

The genus is widespread in the New World, but has not been
recently revised. Of the some fifteen described species, only one
was previously known in postlarval stages. Nymphs and adults of
Hannemania hylae (Ewing) were described by Ewing (1926:266,
figs. 3-4). His descriptions and figures are similar to the species
described here, in general, but he did not give enough details for
specific recognition.

Nymphs of the Trombiculidae 201

Key to Species of Hannemania (Nymphs)

1. Cheliceral blade rather broad (fig. 19) multifemorala p. 203

r. Cheliceral blade narrow (fig. 20) 2

2. Generally smaller; mean ASL about 150 jj.> range 117-170 -x;
mean SB about 34 [x, range 21-38 pt; distance across both eyes

about 40 jj, eltoni p. 202

2'. Generally larger; mean ASL about 218 j;,, range 215-220 [x; mean
SB about 53 pi,, range 49-56 ^,; distance across both eyes about
80 [ji dunni p. 201

Hannemania dunni Sambon

(Fig. 204)

Hannemania dunni Sambon, 1928, Ann. Trop. Med. Parasit., vol. 22, p. 129
{larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1246 (larva).

Description of nymph. — Body: About 1.2 mm. long, color red.
Eyes red in life, distance across both eyes about 80 ]}.. Setae thick
but not expanded, branches as usual; tip of each a recurved hook-
like spine; posterior setae about 70 [jl long. Sternal area about
square, sclerotized, with about 15 setae. Measurements of tarsus
I (1 specimen); TL: 270, TH: 110, TL/TH: 2.5.

Gnathosoma: Basis capituli of normal shape. Cheliceral base of
usual shape, with numerous conspicuous puncta; blade shearlike
but narrowed, with about 23 small teeth, apodeme of blade not
constricted. Cheliceral measurements (means of 3 specimens);
BL: 187.3 ± 4.06, BH: 78.0 ± 3.61, CL: 105.7 ± 2.73, BL/BH:
2.37 ± 0.058, BL/CL: 1.77 ± 0.033. Palpus of usual shape; tarsala
about 18 pi long; tibia without seta at inner tarsal articulation, but
with stiff monopectinate seta below accessory claws; tarsus with 4
inner, 1 outer apical nude setae. Setal counts (1 specimen); femur:
45, genu: 31, tibia: range 7-9, tarsus : range 18-19.

Scutum: Tectal setae apical, set in pits. Apodeme large. Scat-
tered puncta present. Scutal measurements (means of 2 speci-
mens); ASL: 217.5 ± 2.50, SB: 52.5 dr 3.501, TS: 56.0 ± 0.000,
ASL/SB: 4.15 ± 0.2500.

Remarks. — This species, like H. eltoni, proved easy to culture and
as a result more adults were available for study than nymphs. Both
nymphs and adults ate collembola eggs.

Hannemania dunni is very similar to H. eltoni. The characters
used to separate these two as nymphs involve size, dunni being much
the larger.

Loomis (1956:1246) identifies larvae from which these nymphs

202 The University Science Bulletin

were reared, as Hannemania diinni and I accept his identification.
However, these larvae are somewhat larger than other clunni of
Loomis and may be specifically distinct.

Specimens examined. — Total 4 nymphs, as follows: Arkansas.
Montgomery County: Plethodon caddoends, January 26, 1952, taken
by H. A. Dundee (4 nymphs, reared DAC).

Hannemania eltoni Sambon
(Figs. 20, 61,98, 99, 139)

Hannemania eltoni Sambon, 1928, Ann. Trop. Med. Parasit., vol. 22, p. 129
(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1243 (larva).

Description of nymph. — Resembles Hannemania dttnni, except as
follows :

Body: About 0.9 mm. long. Distance across both eyes about 40
a. Setae with branches longer; posterior setae about 45 pi, long.
Measurements of tarsus I (means of 9 specimens); TL: 184.5 ±
6,01, TH: 86.2 ± 2.72, TL/TH: 2.14 ± 0.0503.

Gnathosoma: Basis capituli short, broad. Cheliceral base not
heavily punctate; blade with about 14-29 teeth. Cheliceral measure-
ments (means of 9 specimens); BL: 120.0 zt 5.21, BH: 53.3 ± 2.69,
CL: 74.4 ± 3.00, BL/BH: 2.33 ± 0.110, BL/CL: 1.61 ± 0.035.
Palpus with tarsala about 16 a long. Setal counts; femur: 30.56 ±
2.17 (8 specimens), genu: 18.45 ± 0:575 (10 specimens), tibia:
range 5-8, tarsus: range 10-16.

Scutum: Measurements (means of 9 specimens); ASL: 149,6 zh
5.68, SB: 33.6 d= 1.733, TS: 33.5 ± 0.707, SENS: about 100-150,
ASL/SB: 4.5 ± 0.247.

Remarks. — This species is readily cultured. Both nymphs and
adults eat collembola eggs.

Loomis (1956:1243) discusses the application of the name eltoni
to larvae from which these nymphs were reared.

Specimens examined. — Total 24 nymphs, as follows: Texas.
Bexar County: Camp Bullis, Rana pipiens, April 25, 1954 (8
nymphs, reared RBL). Kansas. Seward County: 12 mi. NE
Liberal, Rana pipiens, September 9-10, 1948 (11 nymphs, reared
LJL). Cheyenne County: 4 mi. N St. Francis, Rana pipiens, July
22, 1948 (3 nymphs, reared LJL). Johnson County: Sunflower,
Acris gryllus, March 29, 1949 (2 nymphs, reared LJL),

Nymphs of the Trombicxjlidae 203

Hannemania multifemorala Loomis

(Figs. 19, 178)

Hannemania multifemorala Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p.
1247 (larva).

Description of nymph. — Resembles Hannemania dunni except as
follows:

Body: About 1.1 mm. long. Eyes present but indistinct. Poste-
rior body setae about 40 [;. long. Measurements of tarsus I ( 1 speci-
men); TL: 170, TH: 75, TL/TH: 2.3.

Gnathosoma: Cheliceral base not heavily punctate; blade rather
broad, upper margin straight, with about 16 small teeth. Cheliceral
measurements (means of 2 specimens); BL: 110.5 ± 10.5, BH:
53.3 zh 4.5, CL: 60.5 ± 9.50, BL/BH: 2.05 ± 0.05, BL/CL: 1.85
± 0.150. Palpus with tarsala about 11 \). long. Setal counts (1
specimen); genu: 25, tarsus: 13 (other segments obscured).

Scutum: Measurements (1 specimen); ASL: 118, SB: 32, ASL/
SB: 3.7.

Remarks. — The presence of this species, which occurs with H.
eltoni, was not detected until the latter part of our work. It does
not culture well, and cultures of Hannemania from Kansas appear
to be pure eltoni. Possibly culture conditions favorable to eltoni
eliminate multifemorala.

Specimens examined. — Total 3 nymphs, as follows: Texas. Erath
County: Stephenville State Park, Microhyla olivacea, April, 1952,
taken by H. S. Fitch (1 nymph, reared DAC), Nebraska. Rich-
ardson County: ]2 mi. W Verdon, Rana pipiens, August 20, 1948 (2
nymphs, reared LJL).

Genus Whartonia Ewing

Whartonia Ewing, 1944, Proc. Biol. Soc. Washington, vol. 57, p. 102 (type
Hannemania nudosetosa Wharton); Audy, 1954, Stud. Inst. Med. Res. Fed.
Malaya, no. 26, p. 164 (nymph).

Diagnosis (nymph). — (1) Body shape oval, not constricted. (2)
Posterior body setae 7nodified, leaflike. (3) Ratio TL/TH: 2.4.
(4) Tarsus I rather pyriform.

Additional features (nymph). — (5) Body size medium (0.8 mm.
long). (6) Eyes absent. (7) Adjacent body setae of similar
lengths. (8) Sclerotized sternal plate present. (9) Legs not un-
usually elongated; branched setae slightly expanded. (10) Chelic-

204 The University Science Bulletin

eral blade narrowed, nearly shearlike. ( 11 ) Palpus stout, moderate
in size; trochanter short, broad; some femoral, some genual setae ex-
panded; claw moderate in size; tarsus with seven branched setae
and five apical nude setae. ( 12) Hypostome blunt, with six to eight
apical setae with reduced branching; about ten branched, slightly
expanded ventral setae. ( 13 ) Basis capituli of normal shape. ( 14 )
Tectum large, well-sclerotized, saggitate in shape; tectal setae
forked, not expanded. (15) Crista a narrow rod. (16) Sensillary
area oval; ratio ASL/SB: 3.6. (17) About ten to fifteen parascutal
setae.

Remarks. — The above characterization is based upon Acoma-
tacarus senase Greenberg, which Loomis (1956:1241) transferred
to Whartonia. Although this species does fit the generic diagnosis
given by Wharton and Fuller (1952:104) for larvae, possibly it is
not congeneric with Whartonia nudosetosa, the genotype.

Morphologically Whartonia senase is quite similar to Acoma-
tacarus. Some features appear to be of group importance, includ-
ing the narrowed cheliceral blade (10), the short palpal trochanter
(11), the blunt hypostome (12) and the forked tectal setae (14).
The leaflike modification of the posterior body setae is the most
obvious character for recognition.

Besides W. senase, four species of Whartonia have been described
and while some have evidently been reared, the nymphal descrip-
tions have not been published.

Whartonia senase (Greenberg)
(Figs. 36, 60, 94, 95, 143, 182, 187, 234)

Acomatacarus senase Greenberg, 1952, Ann. Ent. Soc. America, vol. 45, p. 484

( larva ) .
Whartonia senase, Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1242

( larva ) .

Description of nymph. — Body: About 0.8 mm. long, color not
known. Setae modified as leaflike strvictures; central stem of each
evident but bearing two lateral flanges with sclerotized margins and
weblike surfaces; margins of flanges irregular; central stem with
small branches; posterior setae about 25 \). long. Sternal area con-
taining broad sclerotized plate, with 8 expanded setae. Measure-
ments of tarsus I (1 specimen) TL: 141, TH: 60, TL/TH: 2.4.

Gnathosoma: Cheliceral base stout, punctate, area of articula-
tion with basis capituli slightly shortened; blade rather narrow but
curved, about 10 tiny teeth on apical portion, apodeme of blade not
constricted. Cheliceral measurements (1 specimen); BL: 60, BH:

Nymphs of the Trombiculidae 205

34, CL: 38, BL/BH: 1.8, BL/CL: 1.6. Palpus with dorsal femoral
and dorsal genual setae slightly expanded; claw slender; tarsala
about 8 [L long; tarsus with 3 inner, 2 outer apical nude setae. Setal
counts (1 specimen): femur: 12, genu: 10.5, tibia: range 4-5, tar-
sus, 7.

Scutum: Tectum with margin conspicuously sclerotized; setae
forked at about half their length. Crista broadened gradually
towards posterior. Apodeme short, blunt. Scutal measurements
(1 specimen); ASL: 107, SB: 30, TS: 25, SENS: (obscured),
ASL/SB: 3.6.

Specimens examined. — Total one, as follows: Kansas. Barber
County: S'A mi. S, 1 mi. W Aetna, Myotis velifer, April 10, 1949 ( 1
nymph, reared LJL ) .

Subfamily Trombiculinae Ewing
Ewing, 1929, Manual of External Parasites, p. 22.

Diagnosis (nymph). — Body constricted, figue 8-shaped. Tectum
with one or no setae. Sensilla variable, with at least short basal
branches (except Blankaartia). Anus obviously divided into two
shoelike portions. Setae on palpal genu more numerous than those
on palpal femur, or about the same number on these segments.

Additionnl features (nymph). — Eyes usually absent; when pres-
ent 1/1 and either anterior to or adjacent to sensillary area. Ster-
num variable, rarely open behind or with precoxal plates of coxae
I; usually closed behind by a crossbar connecting coxae II. Scutum
with tectal margin variable; mean ratio ASL/SB: about 1.0-3.2;
bulla usually prominent, sometimes small; lateral keels present;
carina usually present; ridges often present in sensillary area.
Gnathosoma with cheliceral base elongated to unusually elongated
(mean ratio BL/BH: 1.9-4.0); articulation of cheliceral base with
basis capituli usually ventral but sometimes terminal; posterior tip
of cheliceral base elevated or depressed. Palpus usually with t\vo
accessary claws, rarely only one or more than two. Pharynx usu-
ally inconspicuous but sometimes marked with a reticulate pattern.

Diagnosis (larva). — (Modified from Wharton and Fuller, 1952:
41). Leg I with seven segments, legs II and III with six or seven
segments; anterior, median projection of scutum absent; antero-
median scutal seta present or absent; paired submedian scutal setae
absent.

Included genera. — Known from both larvae and postlarvae:
Ascoschongastia Ewing, Blankaartia Oudemans, Cheladonta Lipov-

206 The University Science Bulletin

sky et al., Doloisia Oudemans, Euschongastia Ewing, Euschongas-
toides Loomis, Guntherana Womersley and Heaslip, Heaslipia
Ewing, Neoschongastia Ewing, Pseudosichongastia Lipovsky, Sclion-
gastia Oudemans, Schoutedenichia Jadin and Vercammen-Grand-
jean, Speleocola Lipovsky, TronibicuJa Berlese, Walchia Ewing,
and Walchiella Fuller, Known only from larvae: Anominalaspis
Brennan, Bahiangia Southcott, Endotrombicula Ewing, Gahrliepia
Oudemans, Gateria Ewing, Giroudia Vercammen-Grandjean, Mac-
kiena Traub and Evans, Mijotrombictda Womersley and Heaslip,
Novotrombicula Womersley and Kohls, Oenoschongastia Womers-
ley and Kohls, Riedlinia Oudemans, Sauracarella Lawrence, Schon-
gastiella Hirst, T ecomatlana Hoffmann and Womersia Wharton.
Known only from adults: Ipotromhicula Womersley and Speotrom-
bicula Ewing,

Remarks. — Of the sixteen genera known only as larvae, nine are
monotypic; only two of sixteen genera known from both larvae and
postlarvae are monotypic. The two genera known only as adults
are monotypic and are obviously based on single characters.

The subfamily Trombiculinae as represented here includes genera
formerly placed in the subfamily Walchiinae ( == Gahrliepiinae ) .

As postlarvae the genera are divisible into two groups. The
genera Euschdngastoides, Pseudoschongastia, and Walchia (and
probably Schoutedenichia) share the following features: Cheliceral
base with posterior tip elevated, area of articulation with basis
capituli ventral; hypostome short, blunt, with about twenty short
apical nude or nearly nude setae; pharynx marked with a reticulate
pattern; scutum short and broad, mean ratio ASL/SB: about 1.2-1.7.

Remaining genera have the cheliceral base with posterior tip de-
pressed, area of articulation with basis capituli ventral; hypostome
elongated, tip usually pointed but sometimes blunt, with about eight
apical nude setae; pharynx not marked with a reticulate pattern;
scutum longer, mean ratio ASL/SB: 2.0 or greater (except Gunther-
ana and Doloisia). However, the use of formal names for these
groups would be premature.

Species studied by the writer are members of the genera
Blankoartia, CheJadonta, Euschongastia, Euschdngastoides, Neo-
schongastia, Pseudoschongastia, Speleocola, Trombicula, and Wal-
chia. Descriptions of other postlarvae are sufficiently complete so
that the remaining genera can be keyed out, except for Ascoschon-
gastia and Walchiella, and several subgenera. In the following dis-
cussions the genera seen by the writer will be presented first, and
those not seen will follow.

Nymphs of the TROMBicuLroAE 207

Key to Genera of Trombiculinae (Nymphs)

1. Leg I with a pair of distally trifurcate claws. . . . Speotrombicula* p. 274
1'. Leg I with a pair of distally undivided claws 2

2. Scutum with a branched body seta at the base of the crista, in

the sensillary area Ipotromhicula* p. 273

2'. Scutum without a body seta in the sensillary area 3

3. Precoxal plates of coxa I present and separated, not fused in the
midline to form a longitudinally divided sternum 4

3'. Precoxal plates of coxa I usually absent; if present they are
fused in the midline to form a longitudinally divided sternum .... 6

4. Long and short body setae intermixed (American species with
extremely elongated palpal claws) Neoschongastia p. 231

4'. Adjacent body setae of similar lengths; palpal claws not un-
usually elongated 5

5. Sensillary area narrower (ratio ASL/SB: about 2 in nymphs)

Schongastia* p. 274
5'. Sensillary area wide (ratio ASL/SB: about 1 in nymphs)

Guntherana* p. 272

6. Tarsus I vdth a stumplike process on dorsoapical margin 7

6'. Tarsus I without a stumphke process on dorsoapical margin 8

7. Tectal seta absent Walchia p. 268

7'. Tectal seta present Schoutedenickia* p. 274

8. Cheliceral base unusually elongated distally (ratio BL/BH:
about 4.0 in nymphs); cheliceral blades rather short, semilunar;

palpal trochanter elongated Cheladonta p. 211

8'. Cheliceral base only rarely unusually elongated (ratio BL/BH:
up to 3.6 in nymphs); cheliceral blades longer, shearlike; palpal
trochanter not elongated 9

9. Hypostome short, blunt, with about twenty short apical nude or
nearly nude setae; pharynx marked with a reticulate pattern ... 10

9'. Hypostome long, pointed, with about eight longer apical nude
setae; pharynx indistinct 11

10. Tectal margin without serrations; sensilla stiff, thick to subclavate;

mean ratio ASL/SB: about 1.7 Pseuduschongastia p. 235

10'. Tectal margin with serrations; sensilla almost flagelliform, flexi-
ble; mean ratio ASL/SB: about 1.3 Euschongastoides p. 226

11. Eyes present in front of and separated from sensillary area; sen-
silla completely nude Blankaartia p. 208

ir Eyes usually absent; when present, at the sides of and adjacent
to sensillary area; sensilla with at least short basal branches. ... 12

12. Tectal margin without teeth or serrations 13

12'. Tectal margin with teeth or serrations 14

13. Tectum large or small but hyaline, not developed into a sclero-

tized "nase" Euschongastia p. 215

13'. Tectum large, an elongated triangular cone, not hyaline, developed

into a conspicuous "nase" Heaslipia* p. 273

14. Posterior body setae short and with compound branching

Speleocola p. 238

* Specimens were not seen by the writer.

208 The University Science Bulletin

14'. Posterior body setae short or long but never with compound

branching 15

15. Sensillary area narrow; mean ratio ASL/SB: 1.9-2.7 Trombicula p. 258
15'. Sensillary area wide; mean ratio ASL/SB: 0.7-1.2 Dohisia* p. 271

Genus Blankaartia Oiidemans

Bkmkaartia Oudemans, 1911, Ent. Ber., vol. 3, p. 123 (type Trombidium

niloticurn Trdgnrdh, 1904) (adult).
Trombicula (Blankaartia), Fuller and Wharton, 1951, Psyche, vol. 58, p. 87;

Wharton et al, 1951, Jour. Parasit., vol. 37, p. 29; Wharton and Fuller,

1952, Mem. Ent. Soc. Washington, no. 4, p. 42.
Trombicula (Trdgardhula) Berlese, 1912, Redia, vol. 8, p. 4 (type Trombidium

niloticurn Triigardh, 1904).
Trdgordhula, Womersley, 1948, Trans. Roy. Soc. South Australia, vol. 72, p.

83; Womersley, 1952, Rec. South Australian Mus., vol. 10, p. 114 (post-
larvae).
Pentagonella Thor, 1936, Zool. Anz., vol. 114, p. 30 (type Trombidium. ardeae

Triigardh, 1904).
Trojnbicula (Megatrombicida) Michener, 1946, Ann. Ent. Soc. America, vol.

39, p. 432 (type Trombicula alleei Ewing, 1926).

Diagnosis (nijmph). — (1) Eyes 1/1, separated from sensillary
area. (2) Precoxal plates absent. (3) Body setae branched, not
expanded, no compound branching; adjacent setae of similar
lengths. (4) Claws on tarsus I undivided distally. (5) Tarsus I
without dorsoapical stumplike process. (6) Cheliceral base not
unusually elongated; blade shearlike. (7) Palpal trochanter not
elongated. (8) Hypostome pointed, projecting, with about eight
apical nude setae. (9) Pharynx indistinct. (10) Tectum tri-
angular, somewhat sclerotized, with smooth margin; with a single
tectal seta. (11) Sensillary area triangular; mean ratio ASL/SB:
2.9-3.2. (12) Without branched body seta in sensillary area.
(13) Sensilla flagelliform, long, entirely nude.

Additional features (nymph). — (14) Body size medium (about
0.8-0.9 mm. long), of usual shape. (15) Sternum pentagonal,
closed behind. (16) Tarsus I elongated. (17) Articulating area
of cheliceral base with basis capitiili ventral; base with posterior
tip depressed. (18) Palpus stout, large; distal portion of tibia
unusually elongated; palpal claw short, slightly curved; two to
seven accessory claws; tarsus with about seven branched setae and
four or five apical nude setae. ( 19 ) Hypostome with about tsventy
ventral branched setae. (20) Basis capituh of usual shape. (21)
Crista narrow, elongated. (22) Sensillary area with basal plate of
two bandlike halves; bulla elongated anteriorly; carina apparently
absent; sensillary bases placed well lateral; anterior, lateral, and
posterior ridges somewhat weak; ridges medial to sensillary bases
present; apodeme normal. (23) About thirty parascutal setae.

* Specimens were not seen by the writer.

Nymphs of the Trombiculidae 209

Remarks. — Fuller and Wharton (1951:85) have discussed the
proper name for this group. I follow them in considering Pentago-
nella and Megatrombicida as synonyms. Womersley has persisted
in using the name Trdgdrdhula.

Previously Blankaartia has been accommodated as a subgenus
of Trombicula (except by Womersley, who uses generic status).
The reasons appear to have been twofold; while Blankaartia post-
larvae appear distinct, little is known of Trombicula postlarvae, and
Blankaartia larvae are very similar to certain Trombicula larvae.
This situation has not changed, except that more Trombicula post-
larvae have become known.

I have decided to follow Womersley in giving generic status to
this group. Nymphs of Blankaartia appear related to those of
Trombicula but are amply distinct. The larvae are more closely
similar but can apparently be separated (by keys to subgenera of
Trombicula as given by several authors).

Distinctive features of Blankaartia appear to be primitive ones
as well. The position of the eyes, the entirely nude sensilla, and
the smooth tectal margin are leeuwenhoekiine characters. The
numerous accessory claws are also found in the Trombidiidae. The
scutum, while triangular, is distinct from that of Trombicula in
possessing an elongated bulla and lacking a carina.

The diagnosis given above is based upon the two species B. alleei
and B. velascoi. Additional postlarvae are known: B. nilotica and
B. peruviana are known only as adults; postlarvae of B. acuscutel-
laris and B. attenuata have also been described. A single species,
B. ardeae, is known only as the larva. Descriptions of known post-
larvae may be found in Womersley, 1952, pages 314-323.

The generic diagnosis given above differs from descriptions of
Womersley in an important respect. Womersley describes the
tectum ("epistome") as bearing "fine denticulations." In specimens
(including adults) of B. alleei, B. attenuata (adults only), and B.
velascoi, examined by me, the tectal margin appears smooth. Wo-
mersley's drawing of B. velascoi shows the tectal margin as nearly
smooth. The tectal margins of B. acuscutellaris and B. nilotica are
illustrated as possessing more marked teeth. This character requires
confirmation.

Womersley includes in Trdgnrdhula (= Blankaartia) the species
Trombicula japonica (Tanaka), on the basis of Tanaka's descrip-
tion of eyes anterior to the sensillary area. However, Sasa (1953:
423) describes the nymph as lacking eyes, and places it in Trom-
bicula (Neotrombicula) , as do Philip and Fuller (1950:50). It is

210 The University Science Bulletin

evident that Womersley is dealing with a different form than is
Sasa. I follow Philip and Fuller, and Sasa, and place japonica in
Tromhicida ( Neotrombicula ) .

Besides B. alleei and B. velascoi, only B. acuscutellaris is known
also as a nymph. Womersley (1952:317) describes the nymph of
acuscutellaris but gives no characters to separate it from alleei.
Michener (1946:434) presents a key which separates nymphs of
alleei and velascoi.

Blankaartia alleei (Ewing) New Combination

(Figs. 22, 63, 171, 172, 208)

Trombicula alleei Ewing, 1926, Ent. News, vol. 37, p. Ill (adult).
Trombictda (Megatrombicula) alleei, Michener, 1946, Ann. Ent. Soc. America,

vol. 39, p. 434 (larva, nymph, adult).
Trdgardhula alleei, Womersley, 1948, Trans. Roy. Soc. South Australia, vol. 72,

p. 89 (adult); Womersley, 1952, Rec. South AustraHan Mus., vol. 10, p. 319

( postlarvae ) .
Trombicula (Blankaartia) alleei, Wharton and Fuller, 1952, Mem. Ent. Soc.

Wasliington, no. 4, p. 43.

Description of nymph. — Body: About 0.9 mm. long, color red.
Eyes about 39 [x in diameter, color not known. Body setae normal,
tips simple; posterior setae about 50 [i long. Sternum with about
10 setae. Measurements of tarsus 1(1 specimen); TL: 224, TH:
117, TL/TH: 1.9.

Gnathosoma: Cheliceral base elongated, punctate; blade elon-
gated but shearlike, nearly devoid of teeth (two tiny teeth present
on one chela of single specimen ) . Cheliceral measurements ( 1
specimen); BL: 170, BH: 68, CL: 104, BL/BH: 2.5, BL/CL: 1.61.
Palpus with dorsal femoral and genual setae with reduced branch-
ing, some nude; tibia with stalklike distal elongation bearing stout
claw, setae nude, 3 accessoiy claws on small dorsal flange; tarsala not
seen; tarsus with but 5 branched setae with reduced branching, 4
apical nude setae, arranged 2 inner, 2 outer. Setal counts ( 1 speci-
men); femur (obscured), genu: 14, tibia: 9, tarsus: 5.

Scutum: Lateral keels not seen. Crista very narrow. Sensillary
area triangular but elongated anteriorly (apex of triangle long);
bands of basal plate narrow; bulla prominent. Scutal measurements
(1 specimen); ASL: 205, SB: 70, TS (obscured), SENS (obscured),
ASL/SB: 2.9.

Remarks. — This description is based upon a single specimen
reared by Dr. C. D. Michener.

Specimens examined. — One nymph, as follows: Panama. Colon
Province: Santa Bosa, Myiozetetcs cajonensis, November 11, 1945
(1 nymph, reared by C. D. Michener).

Nymphs of the Trombiculidae 211

Blankaartia velascoi (Boshell and Kerr), New Combination

(Figs. 148, 169, 170,206)

Trombicula velascoi Boshell and Kerr, 1942, Rev. Acad. Colombiana Cien.

Exact., Fisico-Quim. Y Nat., vol. 5, p. 113 (adult).
Tro7nbicula (Megatrombicula) velascoi, Michener, 1946, Ann. Ent. See. America,

vol. 39, p. 438 (larva, mjmph, adult).
Trdgdrdhula velascoi, Woniersley, 1948, Trans. Roy. Soc. South Australia, vol.

72, p. 89 (adult); Womersley, 1952, Rec. South Austrahan Mus., vol. 10,

p. 321 (postlarvae).
Trombicula (Blankaartia) velascoi, Wharton and Fuller, 1952, Mem. Ent. Soc.

Washington, no. 4, p. 44.

Description of nypmh. — Resembles B. alleei except as follows:

Body: About 0.8 mm. long, color red. Eyes about 32 \i. in diam-
eter. Posterior body setae longer, about 90 [l long; branches short;
tips simple, attenuated. Measurements of tarsus I (means of 4
specimens); TL: 208.0 ± 4.97, TH: 80.5 ± 2.06, TL/TH: 2.58
± 0.0855.

Gnathosoma: Cheliceral blade with 21-30 small teeth (but one
specimen without teeth). Cheliceral measurements (means of 3
specimens); BL: 165.3 ± 3.33, BH: 54.0 ± 0.00, CL: 88.2 ± 4.13,
BL/BH: 3.0 ± 0.00, BL/CL: 1.97 ih 1.14. Palpal tibia with setae
showing reduced branching; 4 to 7 accessory claws; tarsala about
13 [X long; tarsus with 5 apical nude setae arranged 3 inner, 2 outer.
Setal counts (means of 3 specimens); femur: 8.17 dz 1.833, genu:
17.3 ± 2.892, tibia: range 4-7, tarsus: range 6-9.

Scutum: Measurements (means of 4 specimens); ASL: 155.8 ±
2.531, SB: 48.3 ±1.0308, TS: 56 (1 specimen), SENS: about 160,
ASL/SB: 3.23 ±0.0408.

Specimens examined. — Total 6 nymphs, as follows: Panama.
Canal Zone: Juan Mina, Myiozetetes similis, October 3, 1945 (4
nymphs, reared by C. D. Michener); October 30, 1945 (1 nymph,
reared by C. D. Michener). Colon Province: Gatuneillo, Saltator,
October 30, 1945 ( 1 nymph, reared by C. D. Michener).

This material, as well as that of B. alleei, was kindly lent by Dr.
W. V. Gertsch of the American Museum of Natural History.

Genus Cheladonta Lipovsky, Crossley and Loomis

Cheladonta Lipovsky, Crossley and Loomis, 1955, Jour. Kansas Ent. Soc, vol.
28, pp. 137-139 (type Cheladonta niiclieneri Lipovsky, Crossley and
Loomis) (larva).

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates ab-
sent. (3) Body setae slightly expanded, branched, without com-
pound branching; adjacent setae of similar lengths. (4) Claws on
tarsus I undivided distally. (5) Tarsus I without dorsoapical

212 The University Science Bulletin

stumplike process. (6) Cheliceral base unusually elongated (mean
ratio BL/BH: about 4.0), blade curved, semilunar-sJiaped. (7)
Palpal trochanter elongated. ( 8 ) Hypostome unusually elongated,
pointed; with about eight short apical nude setae. (9) Pharynx in-
distinct. (10) Tectum nearly square, hyaline, with or without short
teeth on margin, with one tectal seta. (11) Sendllary area nearly
reniform; mean ratio ASL/SB: about 2.1. (12) Without branched
body seta in sensillary area. (13) Sensilla straight to subclavate,
branches variable.

Additional features (nymph). — (14) Body size medium (about
0.6-0.7 mm. long), elongated. (15) Sternum roughly triangular,
closed behind. (16) Tarsus I rounded. (17) Articulating area of
cheliceral base with basis capituli ventral, small; base with pos-
terior tip depressed. (18) Palpus somewhat elongated; size mod-
erate; distal portion of tibia not unusually elongated; claw mod-
erate in size, slightly curved; two accessory claws; tarsus with
about nine branched setae and one apical nude seta. (19) Hypo-
stome with about ten to twenty ventral setae, nude or nearly so.
(20) Basis capituli of usual shape. (21) Crista not narrowed,
rodlike. (22) Sensillary area with basal plate of two rather broad
bands; bulla reduced in size; carina absent; sensillary bases rather
small, placed well forward; anterior, lateral, and posterior ridges
present but no ridges medial to sensillary bases; apodeme broad
and prominent. (23) About ten parascutal setae.

Remarks. — The genus Cheladonta was erected for the species C
micheneri, C. crossi, and C ouachitensis from the United States, and
Neoschongastia ikaoensis from Japan and Korea. The foregoing
diagnosis is based upon nymphs of micheneri and ouachitensis, and
Sasa's (1953:426) description of the nymph of ikaoensis. Sasa il-
lustrates no teeth on the tectal margin and states that the tectum
is "without conspicuous denticles along anterior margin." Other-
wise, his description agrees closely with the American species.

The characters distinctive for Cheladonta appear to be (6), (7),
(16), (19), the shape of the sensillary area under (11), and the
broad scutal apodeme under (22). In combination these charac-
ters indicate very distinctive forms; Cheladonta postlarvae are im-
mediately recognized as such.

Larvae of Cheladonta seem close to Euschongastia or Neoschon-
gastia but possess a key character in the ventral serrations on the
cheliceral blade. In contrast, postlarvae appear far removed from
other genera. The general appearance of the scutum suggests that

Nymphs of the Trombiculidae 213

of Neoschongastia, and indeed these two may be closely related.
Vercammen-Grandjean (in correspondence) indicates rather close
similarities between larvae of Cheladonta and larvae of certain
species of Schoiitedenichia; however, postlarvae of these two gen-
era are markedly different.

Key to Species of Cheladonta (Nymphs)

1. Sensilla subclavate, greatest width near apex; cheliceral blade with

teeth large, conspicuous micheneri p. 213

r. Sensilla slightly expanded but greatest width in midregion, tapered
distally; cheliceral blade with teeth small to tiny 2

2. Tcctal margin with obvious teeth ouachitensis p. 214

2'. Tectal margin without obvious teeth ikaoensis.

(See remarks on Cheladonta, p. 212.)

Cheladonta micheneri Lipovsky, Crossley and Loomis

(Figs. 42, 43, 85, 183, 184, 195, 196)

Cheladonta micheneri Lipovsky, Crossley and Loomis, 1955, Jour. Kansas Ent.
Soc, vol. 28, p. 137 {larva).

Description of nymph. — Body: About 0.7 mm. long, color orange
to white. Setae short, somewhat expanded, tips rounded; branches
short and fine; posterior setae about 15 \i long. Sternal area roughly
triangular in shape, distinctive; usually closed behind, occasionally
with a trace of longitudinal division; with about 8 setae. Legs
appear shortened. Measurements of tarsus I (means of 9 speci-
mens); TL: 111.1 ± 2.010, TH: 72.2 ±1.847, TL TH: 1.56 ±
0.0294.

Gnathosoma: Cheliceral base peculiarly elongated, somewhat
broadened near proximal end but tapering to apex; puncta few;
blade short, semilunar, with about 11 teeth, mostly large and con-
spicuous; apodeme of blade strongly constricted. Cheliceral meas-
urements (means of 8 specimens); BL: 121.3 ± 3.21, BH: 30.4
± 0.460, CL: 34.0 ± 1.02, BL/BH: 4.0 ± 0.139, BL CL: 3.58 ±
0.084. Palpi rather distinctive in appearance, because of elongated
trochanters and genu wider than long; tarsala about II \t. long;
tarsus with apical nude seta on inner surface. Setal counts (means
of 8 .specimens); femur: 4.44 ± 0.333, genu: 8.25 ± 0.3134, tibia:
range 4-6, tarsus: range 8-10.

Scutum: Sensillary area nearly reinform but anterior projection
present; broad bands of basal plate well separated; apodeme often
with irregular markings, furcate at posterior tip; bulla small, tri-
angular; sensilla subclavate with short fine branches which lengthen
towards tip. Scutal measurements (means of 9 specimens); ASL:
75.1 ± 1.67, SB: 35.4 ± 1.001, TS: 35.3 ±: 0.833, SENS: about 70,
ASL^SB: 2.14 ±0.0377.

214 The University Science Bulletin

Remarks. — In culture, nymphs of C. micheneri ate active stages
of collembolans and refused collembola eggs. As larvae they are
rarely abundant upon hosts and large numbers were not available
for culture purposes.

Specimens exom,ined. — Total 12 nymphs, as follows: Kansas.
Barber Coimtij: 1 mi. W, 4/2 mi. S. Aetna, Cijnomys hidovicianus,
July 27, 1952 (3 nymphs, reared DAC). Douglas County: 3 mi.
W. Lawrence, Sylvilagus floridamis, November 12, 1949 ( 2 nymphs,
reared LJL); Lawrence, ISleotoma foridana nest, FebiTiary 20, 1950
(4 nymphs, reared LJL). Jefferson County: % mi. E. 5/2 mi. N
La\vrence, Peromyscus leucopus, January 26, 1952 ( 1 nymph, reared
DAC ) . Norris County: 2 mi. S Council Grove, Peromyscus manic-
idatus. May 31, 1950 ( 1 nymph, reared LJL ) .

Cheladonta ouachitensis Lipovsky, Crossley and Loomis

(Figs. 10, 44, 155, 200, 221)

Cheladonta ouachitensis Lipovsky, Crossley and Loomis, 1955, Jour. Kansas
Ent. Soc, vol. 28, p. 139 (larva).

Description of nymph. — Resembles C. micheneri except as fol-
lows :

Body: About 0.6 mm. long, color not known. Posterior body setae
about 20 \}. long. Measurements of tarsus I ( means of 3 specimens ) ;
TL: 108.7 ± 2.404, TH: 68.0 ± 2.517, TL/TH: 1.60 ± 0.0578.

Gnathosoma: Cheliceral blade with about 13 small or tiny teeth.
Cheliceral measurements (means of 3 specimens); BL: 109.0 ±:
2.64, BH: 27.3 it 1.333, CL: 30.0 ± 1.15, BL/BH: 4.0 ± 0.116,
BL/CL: 3.67 ± 0.1333. Palpus with tarsala about 8 tJ. long. Setal
counts; femur (2 specimens): 3.5 ih 0.0000, genu (3 specimens):
7:83 ± 0.3342, tibia: range 5-6, tarsus: range 6-8.

Scutum: Sensilla slightly expanded in midregion but tapered
distally, not subclavate; distal branches rather long. Scutal meas-
urements (3 specimens); ASL: (not obtainable), SB: 31.6 ± 1.886,
TS: 40.3 ±4.19, SENS: about 80, ASL/SB: (not obtainable).

Remarks. — These nymphs were reared in a culture of Trombicula
(Neotrombicula). Re-examination of records showed that larvae
of C. ouachitensis were also taken from the host; there is little doubt
that these nymphs are ouachitensis.

This species differs from C. micheneri in having smaller, more
numerous cheliceral teeth and in the nature of the sensilla. Judging
from Sasa's description (1953:426), C. ouachitensis is very similar
to C. ikaoensis.

Nymphs of the Trombiculidae 215

Specimens examined. — Total 3 nymphs, as follows: Arkansas.
Polk County: 2 mi. NE Mena, Crijptotis parva, March 3, 1951
(3 nymphs, reared LJL).

Genus Euschongastia Ewing

Euschongastia Ewing, 1938, Jour. Washington Acad. Sci., vol. 28, p. 293 (type
Schongastia sciuricola Ewing) (= Euschongastia americana Ewing) {larva).

Diagnosis {nymph). — (1) Eyes absent. (2) Precoxal plates ab-
sent. (3) Body setae branched, variable; tips of posterior setae
often with tiny branches, sometimes compound; adjacent setae of
similar lengths. (4) Claws on tarsus I undivided distally. (5) Tar-
sus I without dorsoapical stumplike process. (6) Cheliceral base
not unusually elongated; blade shearlike. ( 7 ) Palpal trochanter not
elongated. (8) Hypostome pointed, projecting; with about eight
apical nude setae. ( 9 ) Pharynx indistinct. ( 10 ) Tectum triangular,
hyaline, with margin smooth; with one tectal seta. (11) Sensillary
area triangular to transverse-oval in shape; mean ratio ASL/SB:
2.1-2.7. (12) Without branched body seta in sensillary area. (13)
Sensilla thick, somewhat straight but not subclavate; branches short.

Additional features {nymph). — (14) Body size small to large
(0.4-1.0 mm. long), of usual shape. (15) Sternum closed behind,
roughly pentagonal to elongated in shape. (16) Tarsus I usually
elongated, sometimes nearly rounded. ( 17 ) Articulating area of
cheliceral base with basis capituli ventral; base with posterior tip
depressed. (18) Palpus usually stout, moderate in size, slightly
curved; two accessory claws; tarsus with about nine branched setae
and four or five apical nude setae. ( 19 ) Hypostome with about ten
to twenty ventral branched setae. (20) Basis capituli of usual shape,
rarely broadened. (21) Crista a narrow rod. (22) Sensillary area
with basal plate of two halves; bulla present, sometimes reduced;
carina present; sensillary bases placed well lateral; anterior, lateral,
and posterior ridges present but often weak; ridges medial to
sensillary bases weak; toofhlike projections often present at front
angles of sensillary area; apodeme usually reduced. (23) About
eight to fourteen parascutal setae.

Remarks. — This description is based entirely on the species
studied by the writer. Other postlarvae have been described.
Schongastia samoaensis Womersley and Microtrombidium toestralen-
sis Womersley, both known only as adults, were placed in Schon-
gastia { Ascoschongastia ) ( = Euschongastia ) by Womersley ( 1952 :

216 The University Science Bulletin

423, 383). Also, Womersley described reared nymphs of Neo-
schongastia lanius Radford, Schongastia auchji Womersley, N. kohlsi
Radford, N. mutahilis Gater, and S. nadchairami Womersley and
placed them in Schongastia (Ascoschotigastia). Wharton (1946:
159, 162) gives excellent descriptions of the nymph and adult of
Etischongastia indica (Hirst). More recently, Domrow (1955:57,
130) has described the nymphs of Etischongastia smithi (Womers-
ley) and E. perameles (Womersley). Tromhicula algerica Andre,
known only as adult, was placed in Euschongastia by Wharton and
Fuller (1952:73).

The species known only as adult could as well be accommodated
in Trombicula as in Euschongastia but their final placement will
require more careful descriptions. The species known as both
larva and nymph must be more carefully considered.

The only useful character for separating nymphs of Euschongastia
from those of Trombicula is the nature of the tectum. In Euschon-
gastia, the tectal margin is smooth; in Tromhicula, the tectal margin
has serrations or teeth. In the larval stage a single easily dis-
cernible character separates these two genera; Euschongastia has
expanded sensilla while Tromhicula has flagelliform sensilla. Other-
wise, the two genera appear similar in both stages. The nymphs
of species of Euschongastia as described by Domrow, Wharton,
and Womersley all possess serrations on the tectal margin and
thus in that key couplet would fall into Troinbicula. The larvae,
however, have expanded sensilla and so would be placed in
Euschongastia.

Three of these nymphs, Euschongastia indica, E. perameles, and
E. smithi, would apparently key out to Euschongast aides rather
than Tromhicula or Euschongastia; these species are further dis-
cussed under Euschongastoides. The remainder, as described by
Womersley, would seemingly key to Tromhicula. In the illustra-
tions of these species, the tectum is shown as rounded and with
fine serrations. Possibly these are in reality absent. Also, Womers-
ley's nymphs are Asiatic while mine are American; possibly there
are real group differences between them.

The genus Euschongastia contains a broad assemblage of species
and is long overdue for revision and probable recognition of other
genera for some species groups. Also, its relations with Tromhicula
must be critically surveyed. Any generic separation based upon a
single character is likely to be artificial.

Nymphs of the Trombiculidae 217

Key to Species of Euschongastia ( Nymphs )

1. Posterior body setae ending in single long, fine, attenuated tips,

never with minute branches near tips pipistrelli p. 222

r. Posterior body setae ending in one or (usually) more small or mi-
nute branches 2

2. Sensilla with apical portions nude and attenuated, or with short
branches very sparse on apical portions 3

2'. Sensilla vdth short branches along entire length 4

3. Sternal area elongated-rectangular joned p. 220

3'. Sternal area nearly pentagonal, not elongated posteriorly, trigenuala p. 225

4. One or two rather inconspicuous toothUke projections on anterior
margin of sensillary area directly in front of sensillary bases 5

4'. Three or four prominent, broad toothlike projections arising from
most of anterior margin of sensillary area 7

5. Larger species, ASL about 120 jx> TS about 50 [/,, BL about

135 [X setosa p. 223

5'. Smaller species; ASL about 80-90 ^, TS about 25-30 ^, BL about
100 [X 6

6. Posterior body setae ending in blunt tips with one or two short,
minute, thick branches cynomyicola p. 218

6'. Posterior body setae each ending in attenuated tip divided into

two to four short, attenuate branches criceticola p. 217

7. Tips of posterior body setae shghtly enlarged and ending in numer-
ous short minute compound branches; one (rarely two) lateral
branch at each tip characteristically enlarged and prominent

peromysci p. 221
7'. Tips of posterior body setae not enlarged, ending in two or three
short, often compound branches; without such a modified lateral
branch at tips diversa p. 219

Euschongastia criceticola Brennan
(Figs. 46, 70, 130, 131, 150, 217)

Euschongastia criceticola Brennan, 1949, Jour. Parasit., vol. 34, p. 473 (larva);
Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1340 (larva).

Description of nymph. — Body: About 0.7 mm. long, color yellow
to orange. Body setae branched as usual; tips of posterior setae
ending in 2 to 4 short attenuated branches; posterior setae about
60 [). long. Sternum roughly pentagonal, with about 8 setae. Meas-
urements of tarsus I (means of 8 specimens); TL: 118.1 ± 1.86,
TH: 51.0 ± 1.53, TL/TH: 2.34 ± 0.0962.

Gnathosoma: Basis capituli of usual shape. Cheliceral base
somewhat stout, pimcta few; blade stout, shearhke, with about 13
teeth of varying sizes. Cheliceral measurements (means of 10
specimens); BL: 100.7 ± 1.764, BH: 37.5 ±1.241, CL: 46.8 ±
0.554, BL/BH: 2.73 ± 0.120, BL/CL: 2.14 ± 0.043. Palpus normal,
rather small; tarsala about 11 [x long; tarsus with 3 inner, 1 outer

218 The University Science Bulletin

apical nude setae. Setal counts; femur (6 specimens): 3:67 ±
0.3587, genu (7 specimens) : 7.43 ± 0.5279, tibia: range 4-5, tarsus:
range 8-9.

Scutum: Tectum moderate in size. Sensillary area with basal
plate of two narrow bands; apodeme much reduced; bulla triangu-
lar; carina thin; ridges weak; anterior margin often with one or
two small toothlike projections directly anterior to sensillary bases;
sensilla slightly thickened but flexible, with short branches along
entire length. Scutal measurements (means of 7 specimens); ASL:
81.6 ±2.861, SB: 36.8 ± 1.031, TS: 27.0 ± 1.448, SENS: about
100, ASL/SB: 2.14 ± 0.0645.

Remarks. — Etischdngastia criceticola appears to be a widespread
chigger in western North America (see Loomis, 1956:1341). There
may be more than one form of larva considered under this name.
The nymphs described here are assoicated with the larval form
treated by Loomis (1956:1340).

This species was reared on three occasions. There are no rec-
ords of feeding by the nymphs, but since some adults were ob-
tained the nymphs evidently fed. The food was probably collem-
bola eggs wliich were added to the cultures as food material.

Speciinens examined. — Total 41 nymphs, as follows: Kansas.
Barber County: S^-i mi. S, 1 mi. W Aetna, Neotoma micropiis, April
11, 1949 (39 nymphs, reared LJL); 4 mi. S Aetna, Neotoma micro-
pus, October 6, 1951 (1 nymph, reared LJL). Russell County: 5
mi. N, 2 mi. E Graham, Peromyscus maniculatiis, April 27, 1952 (1
nymph, reared DAG).

Euschongastia cijnomyicola Grossley and Lipovsky

(Figs. 50, 67, 212)

Euschongastia cijnomyicola Crossley and Lipovsky, 1954, Proc. Ent. Soc, Wash-
ington, vol. 46, pp. 240-243 {larva); Loomis, 1956, Univ. Kansas Sci. Bull,
vol. 37, p. 1342 {larva).

Description of nymph. — Resembles E. criceticola except as fol-
lows :

Body: About 0.4 mm. long, color not known. Tips of posterior
body setae blunt, ending in one or two minute, but thick, branches;
posterior setae about 35 pi. long. Sternum with about 6 setae. Meas-
urements of tarsus I (means of 10 specimens); TL: 113.0 ± 5.34,
TH: 53.3 ± 1.044, TL/TH: 2.11 ± 0.0722.

Gnathosoma: Gheliceral base stout; blade normal, shearlike,
with about 18 small to tiny teeth. Gheliceral measurements (means
of 10 specimens); BL: 101.6 ± 2.945, BH: 41.5 ± 0.969, GL: 48.9

Nymphs of the Trombiculidae 219

± 1.197, BL/BH: 2.46 ± 0.0748, BL/CL: 2.08 ±: 0.029. Palpus
rather stout, tarsala about 9 [). long. Setal counts (means of 11
specimens); femur: 8.45 ± 0.3763, genu: 9.85 ± 0.6012.

Scutum: Measurements (means of 10 specimens); ASL: 86.8 ±
3.339, SB: 42.4 ± 0.897, TS: 29.6 ± 0.653, SENS: about 110,
ASL/SB: 2.05 + 0.0521.

Remarks. — This species is very similar to, although somewhat
smaller than, Etischongastia criceticola. As larvae the two are
readily separable on several characters. Eiischongastia cynomyi-
cola was cultured once. No records of feeding are available, and
no adults were obtained from the culture. Evidently the nymphs
either refused the collembolans and their eggs, or these were in-
adequate food materials.

Specimens examined. — Total 12 nymphs, as follows: Nebraska.
Hitchcock County: 4 mi. E Stratton, Cynomys hidovicianus, August
8, 1949 (12 nymphs, reared LJL).

Euschongastia diversa Loomis

(Figs. 49, 215)

Euschongastia diversa Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1337
{larva).

Description of nymph. — Body: About 0.6 mm. long, color orange
to yellow. Body setae resembling those of E. criceticola but pos-
terior setae ending in 2 to 4 short branches, these sometimes com-
pound; posterior setae about 50 pi long. Sternum roughly pentag-
onal, with about 10 setae. Measurements of tarsus I (means of 6
specimens); TL: 124.0 ± 3.85, TH: 66.3 ± 1.41, TL/TH: 1.87
It 0.0212.

Gnathosoma: Cheliceral base stout, puncta numerous; blade
stout, shortened, shearlike, with about 13 large to small teeth.
Cheliceral measurements (means of 6 specimens); BL: 102.0
± 3.507, BH: 43.3 ± 1.282, CL:^ 52.3 ± 1.308, BL/BH: 2.38 ±
0.1046, BL/CL: 2.00 ± 0.0516. Palpus somewhat stout, accessory
claws somewhat shortened; tarsala about 10 [i long; tarsus with 3
inner, 1 outer apical nude setae. Setal counts; femur (3 speci-
mens); 5.0 ± 0.000, genu (4 specimens): 8.75 ± 0.595, tibia: range
4-6, tarsus: range 9-10.

Scutum. Tectum moderate in size, seta elongate. Sensillary area
with basal plate of two widely separated bandlike halves; apodeme
much reduced or absent; bulla small, triangular; carina prominent,
often doubled posteriorly; strong ridges anterior to and posterior to
sensillary bases, other ridges weak; anterior margin of sensillary

220 The University Science Bulletin

area with three or four broad, prominent, toothhke projections cov-
ering most of margin; sensillum hke that of E. criceticola. Scutal
measurements (means of 6 specimens); ASL: 95.8 zb 4.489, SB:
36.3 ±1.606, TS: 30.0 ± 0.894, SENS: about 125, ASL/SB: 2.65
± 0.1335.

Remarks. — As both larva and posthirva, E. diversa is very similar
to E. peromysci. Nymphs of these two species are separable by
the nature of the tips of the fjosterior body setae, as indicated in
the key to species. This character varies in both species. Many
specimens of £. diversa have the tips of these setae divided into
several short attenuate branches, but others have these branches
compoundly branched. When a lateral branch arises very near
the tip, the setae resemble those of E. peromysci. In none of the
material examined was there any difficulty in separating these two
species by this character, however.

This species was reared on several occasions. The nymphs uti-
lized collembolan eggs as food, and several adults were obtained.

Specimens examined. — Total 87 nymphs, as follows Kansas.
Douglas County: Lawrence, Neotoma floridana, December 9, 1949
(26 nymphs, reared LJL); 3 mi. W Lawrence, Neotoma floridana,
March 2, 1949 (1 nymph, reared LJL) and March 5, 1949 (53
nymphs, reared LJL); 4 mi. N, 1 mi. E Lawrence, Neotoma flori-
dana, December 9, 1948 (1 nymph, reared LJL); 2 mi. S, 2 mi. W
Pleasant Grove, Neotoma floridana, March 4, 1949 (2 nymphs,
reared LJL). Johnson County: 2 mi. W, 1 mi. N Lenexa, SylviJagus
floridanus, November 18, 1953 (3 nymphs, reared RBL). Miami
County: 2 mi. W, 1 mi. S Louisburg, Sylvilagus floridanus, Novem-
ber 24, 1953 ( 1 nymph, reared RBL, idet. by larval skin ) .

Euschongastia jonesi Lipo\'sky and Loomis

(Figs. 26, 64, 108, 109, 153, 216)

Euschongastia jonesi Lipovskv and Loomis, 1954, Joiir. Parasit., vol. 40, pp.
407-410 (larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1332-
1333 (larva).

Description of nymph. — Body: About 0.7 mm. long, color yellow
to white. Body setae like those of E. criceticola, but posterior setae
ending in tips of 2 to 4 short branches, which are surpassed by a
long semiterminal lateral branch; posterior setae about 90 ijl long.
Sternum elongated-rectangular, with about 8 setae. Measurements
of tarsus I (means of 7 specimens); TL: 147.0 ± 4.999, TH: 71.3
± 1.340, TL/TH: 2.06 ± 0.0429.

Gnathosoma: Basis capituli rather short, broad. Cheliceral base
somewhat stout, puncta numerous; blade shearlike but thinner, and

Nymphs of the Trombicxilidae 221

elongated, with about 20 small to tiny teeth. Cheliceral measure-
ments (means of 7 specimens); BL: 127.3 ± 4.279, BH: 50.0 ±
1.309, CL: 68.0 ± 1.448, BL/BH: 2.56 ± 0.2243, BL/CL: 1.87
± 0.3394. Palpus normal, rather large; tarsala about 15 (x long;
tarsus with 4 or 5 apical nude setae. Setal counts (means of 7
specimens); femur: 4.5 ± 0.2886, genu: 11.64 ± 0.8360, tibia:
range 4-6, tarsus: range 8-10.

Scutum: Tectum moderate in size; seta elongated. Sensillary
area with basal plate normal, apodeme much reduced; bulla tri-
angular; carina prominent; strong ridges anterior and posterior to
sensillary bases; sensilla slightly thickened but flexible; short basal
branches numerous but branches on distal portion few, and short.
Scutal measurements (means of 7 specimens); ASL: 128.5 ± 2.79,
SB: 43.6 ±0.922, TS: 47.0 ± 1.964, SENS: about 150, ASL/SB:
2.67 ± 0.0334.

Remarks. — Etischongastia jonesi is similar to E. pipistreUi par-
ticularly in details of the scutum. The two species are separable
by characters of the gnathosoma but most readily by the nature of
the tips of the posterior body setae: E. jonesi has setae ending in
several short branches, while E. pipistreUi has setae ending in a
single long branch. The two species are also similar as larvae.

The nymphs were readily maintained in culture. They fed upon
collembola eggs.

Specimens examined. — Total 7 nymphs, as follows: Kansas. Bar-
ber Counttj: 4 mi. S Aetna, Peromyscus maniculatus, October 7,
1951 (1 nymph, reared DAC). Cowley County: 2 mi. E Rock,
Feromyscus leucopus, March 7, 1953 (6 nymphs, reared DAC, idet.
by larval skins; reared from larvae associated with the type series).

Etischongastia peromysci (Ewing)
(Figs. 48, 66, 104, 105, 152, 214)

Schdngastia peromysci Ewing, 1929, Ent. News, vol. 40, p. 296 (larva).
Euschongastia peromysci Fuller, 1948, Bull. Brooklyn Ent. Soc., vol. 43, p. 108

(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1334-1335

( larva ) .

Description of nynij)]i. — Resembles E. diversa except as follows:
Body: Posterior body setae ending in slightly expanded tips,
which give rise to several short compound branches, each seta with
a thickened subterminal lateral branch surpassing the tip; posterior
setae about 35 [jl long. Sternum with about 6 setae. Measure-
ments of tarsus I (means of 9 specimens); TL: 133.1 ± 0.588, TH:
73.1 ± 0.7536, TL/TH: 1.83 ± 0.0167.

222 The University Science Bulletin

Gnathosoma: Cheliceral blade with about 18 rather small to tiny
teeth. Cheliceral measurements (means of 9 specimens); BL: 115.4
± 4.176, BH: 48.0 ± 0.957, CL: 60.3 ± 0.965, BL/BH: 2.41 ±
0.0588, BL/CL: 1.88 ± 0.325. Palpus with tarsala about 12 [x long.
Setal counts (means of 9 specimens); femur: 3.83 ± 0.2041, genu:
10.0 ± 0.4961, tibia: range 4-5.

Scutum: Measurements (means of 10 specimens); ASL: 105.9 ±
1.645, SB: 43.4 ± 0.499, TS: 38.6 ± 1.335, SENS: about 145, ASL/
SB: 2.44 ±0.0445.

Remarks. — This species shows the most extreme modification of
the tips of the posterior body setae. The minute compound
branches of the tip are often distributed along the prominent lateral
branch, for about half of its length. The characteristics of these
setae are adequate for recognition of the species. A specimen from
Shelby County, Tennessee, shows a deviate condition: Two in-
stead of one prominent subterminal branches, forming about a
ninety-degree angle. Larvae from which this nymph was reared
are typical peromysci. Since E. peromijsci is primarily an eastern
species, this two-spined tip may be more characteristic.

In culture, nymphs of E. peromysci ate collembolan eggs.

Specimens examined. — Total 19 nymphs, as follows: Kansas.
Cowley County: 2 mi. E Rock, Peromyscus leucopus, March 7, 1953
(7 nymphs, reared DAC, idet. by larval skins). Douglas County:
Lawrence, Sylvilagiis floridanus, Lawrence, November 12, 1949 (1
nymph, reared LJL), and Neotoma floridana, March 30, 1949 (2
nymphs, reared LJL); 5 mi. N, 1 mi. E Lawrence, Neotonui flori-
dana, February 18, 1950 (1 nymph, reared LJL); 2 mi. S, 2 mi. W
Pleasant Grove, Neotoma floridana, March 4 and 5, 1949 (3 nymphs,
reared LJL). Jefferson County: 5/2 mi. N Lawrence, Peromyscus
leucopus, January 25, 1952 (2 nymphs, reared DAC). Johnson
County: Roeland Park, Sylvilagiis floridanus, November 10, 1953 ( 2
nymphs, reared RBL). Tennessee. Shelby County: Peromyscus
leucopus, January 31, 1954 (1 nymph, reared DAC, idet. by larval
skin ) .

Euschongastia pipistrelli Brennan
(Figs. 28, 225)

Euschongastia pipistrelli Brennan, 1947, Jour. Parasit., vol. 33, p. 249 (larva);
Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1331-1332 (larva).

Description of nymph. — Resembles E. jonesi, except as follows:

Body: About 1.0 mm. long, color not known. Body setae hke

those of E. criceticola, with branches rather long; tips single, long,

attenuated; posterior setae about 105 [). long; posterior setae mark-

Nymphs of the Trombiculidae 223

edly longer than dorsal subposterior setae. Sternum roughly pen-
tagonal, with about 6 setae. Measurements of tarsus I (means of
4 specimens); TL: 196.0 ± 5.788, TH: 81.3 ± 1.109, TL/TH:
2.40 ± 0.410.

Gnathosoma: Cheliceral base apparently without puncta; blade
of usual shearlike shape, with about 25 small to tiny teeth. Chehc-
eral measurements (means of 4 specimens); BL: 129.5 ± 3.97,
BH: 55.0 ± 2.35, CL: 67.5 ± 2.33, BL/BH: 2.38 ±: 0.233, BL/CL:
1.93 ± 0.008. Palpus with tarsala about 13 \). long; tarsus with 3
inner, 1 outer apical nude setae. Setal counts (means of 4 speci-
mens); femur: 7.88 ± 1.0784, genu: 15.75 ± 1.4790, tibia: range
4-7, tarsus: range 10-13.

Scutum: Sensillum resembling that of E. criceticola. Scutal
measurements (means of 3 specimens); ASL: 141.3 ± 2.03, SB:
54.8 ±0.629, TS: 41.0 ± 2.000, ASL/SB: 2.60 ± 0.0577.

Remarks. — Eiischongastia pipistrelli is restricted in the larval
stage to bats, while its closest relative, E. jonesi, occurs on other
mammals as well. It seems logical to expect that the nymphal stage
of a bat parasite, particularly a cave bat, would possess morphologi-
cal features indicating adaptation to a cave habitat. However,
E. pipistrelli resembles other species of Eiischongastia more closely
than does E. jonesi, which can evidently persist in caves or in other
habitats.

The larval forms of both species appear similar to the type of the
genus, E. sciiiricola; Loomis placed them in the subgenus Euschon-
gastia.

Specimens examined. — Total 4 nymphs, as follows: Oklahoma.
Adair County: 5 mi. S. Kansas border (9^2 S Aetna, Kansas),
Pipistrelltis siibflavtis, March 5, 1950 (4 nymphs, reared LJL).

Eiischongastia setosa (Ewing)
(Figs. 27, 29, 65, 102, 103, 156, 213)

Tromhicula setosa Ewing, 1937, Proc. Biol. Soc. Washington, vol. 50, pp.

170-171 (larva).
Euschongastia setosa. Fuller, 1948, Bull. Brooklyn Ent. Soc, vol. 43, p. 103

(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1329 (larva).

Description of nymph. — Body: Large, about 1.0 mm. long, color
pale orange. Body setae like those of E. diversa; short branches
at tip never with compound branching; posterior setae about 55 [x
long. Sternum roughly pentagonal, with about 8 setae. Measure-
ments of tarsus I (means of 5 specimens); TL: 174.0 ± 5.162, TH:
72.8 ±: 2.834, TL/TH: 2.04 ± 0.0610.

224 The University Science Bulletin

Gnathosoma: Cheliceral base of usual shape, punctate; blade
shearlike, with about 22 small teeth. Cheliceral measurements
(means of 9 specimens); BL: 136.1 dr 3.116, BH: 53.2 ± 1.128,
CL: 73.6 ± 1.987, BL/BH: 2.57 ± 0.0527, BL CL: 1.86 ± 0.0176.
Palpus large and stout; tarsala about 14 ii long; tarsus with 4 or 5
apical nude setae. Setal counts; femur ( 6 specimens ) : 18.5 ±
0.4831, genu ( 8 specimens ) : 9.94 ± 0.6371, tibia: range 5-8, tarsus:
range 9-11.

Scutum: Tectum large, seta long. Sensillary area with basal
plate of rather characteristic shape; apodeme reduced; bulla tri-
angular; carina thin but prominent; strong ridges anterior and pos-
terior to sensillary bases; other ridges weak; anterior margin of
sensillary area with 2 (occasionally 1) elongated, rather character-
istic toothlike projections directly in front of sensillary bases; sen-
sillum like that of E. criceticola. Scutal measurements (means of 8
specimens); ASL: 123.0 ± 4.957, SB: 46.2 =i= 1.172, TS: 49.5
± 2.044, SENS: about 135, ASL/SB: 2.69 ± 0.0639.

Remarks. — Of the species considered here, E. setosa seems to be
the most closely related to the type of the genus, E. sciuricola, judg-
ing from their larvae. Nymphs of sciuricola are unknown.

Several characters which separate E. setosa from other Euschon-
gastia species involve size; E. setosa is the largest of these nymphs.
Otherwise some scutal details appear distinctive, as noted above.

In culture, nymphs of E. setosa ate Collembola eggs. Several
adults were obtained from the cultures.

Specimens examined. — Total 22 nymphs, as follows: Arkansas.
Polk County: 2 mi. NE Mena, Cryptotis parva, March 3, 1951 (1
nymph, reared LJL). Kansas. Barber County: 4 mi. S Aetna,
Neotoma micropus, October 6, 1951 ( 1 nymph, reared DAC ) , and
Peromyscus leucopus, October 7, 1951 ( 1 nymph, reared DAC ) ; 3/2
mi. S, 1 mi. W Aetna, Neotoma micropus, April 11, 1949 (4 nymphs,
reared LJL ) ; 5 mi. S Sun City, Neotoma micropus, April 12, 1949 ( 1
nymph, reared LJL). Cowley County: 2 mi. E Rock, Peromyscus
leucopus, March 7, 1953 (1 nymph, reared DAC, idet. by larval
skin). Douglas County: 1 mi. N, 5 mi. E Lawrence, Neotoma flori-
dana, November 23, 1951 (1 nymph, reared DAC). Johnson
County: Roeland Park, Sylvilagus fioridanus, November 10, 1953 (9
nymphs, reared RBL); 2 mi. N, 1 mi. W Lenexa, Sylvilagus fiori-
danus, November 18, 1954 (2 nymphs, reared RBL). Russell
County: 5 mi. N, 2 mi. E Graham, Peromyscus maniculatus, April
27, 1952 ( 1 nymph, reared DAC ) .

Nymphs of the Trombiculidae 225

Euschongastia trigenuala Loomis

(Figs. 45, 68, 120, 121, 151, 192, 220)

Euschongastia trigenuala Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 1343 (larva).

Description of nymph. — Body: About 0.8 mm. long, color white.
Setae somewhat expanded, with fine branches; tips rounded, with 2
or 3 shorter branches; posterior setae markedly longer than dorsal
subposterior setae; posterior setae about 85 [x long. Sternum roughly
pentagonal, with about 6 setae. Measurements of tarsus I (means
of 8 specimens); TL: 119.3 ± 2.058, TH: 56.4 ± 1.252, TL/TH:
2.14 ± 0.0497.

Gnathosoma: Cheliceral base stout, puncta numerous; blade
stout, shearlike, apodeme of blade markedly constricted; with about
13 rather large, flat teeth. Cheliceral measurements (means of 10
specimens); BL: 98.0 ±: 1.361, BH: 41.2 ± 0.743, CL: 53.8 ±
1.104, BL/BH: 2.38 ± 0.0133, BL/CL: 1.81 ± 0.0233. Palpus stout,
claw somewhat thin; tarsala about 14 [l long; tarsus with 3 inner,
2 outer apical nude setae. Setal counts (means of 10 specimens);
femur: 4.45 ± 0.1740, genu: 8.65 dz 0.3078, tibia: range 4-6, tar-
sus: range 9-10.

Scutum: Tectum moderate in size. Sensillary area with basal
plate of two bands as usual; apodeme much reduced; bulla tri^
angular; carina thin; rather strong ridges anterior to and posterior to
sensillary bases; other ridges weak; anterior margin sometimes with
inconspicuous toothlike projections directly in front of sensillary
bases; sensilla slightly thickened but flexible, with branches numer-
ous on basal half but distal one third to one fourth entirely nude,
attenuated. Scutal measurements (means of 10 specimens); ASL:
81.8 ± 1.597, SB: 37.2 ± 0.646, TS: 27.8 ± 1.236, SENS: about
116, ASL/SB: 2.21 ± 0.0407.

Remarks. — The closest relatives of E. trigenuala are not obvious.
As both larva and nymph, the species has several distinctive features
but these are not so striking as to suggest that E. trigenuala is far
removed from the other species. N>Tnphs of trigenuala are easily
recognized by the nude, tapered tip of the sensillum. Loomis
(1956-1344) shows that larvae of E. trigenuala seem to prefer
fossorial mammals, suggesting that the postlarvae may live in the
subterranean burrows. No nymphs were reared from truly fossorial
hosts.

Specimens examined. — Total 17 nymphs, as follows: Kansas.
Douglas County: 1 mi. E, 5 m. N Lawrence, Microtus ochrogaster,

8—8920

226 The University Science Bulletin

February 20, 1952 (4 nymphs, reared DAC). Johnson County: 2
mi. N, 1 mi. W Lenexa, Microtus ochrogasier, April 2, 1954 (1
nymph, reared DAC). Lijon County: 2 mi. S Chalk, Perognathus
hispidus. May 31, 1950 (10 nymphs, reared LJL). Russell County:
9 mi. S Russell, Peromyscus maniculatus, April 26, 1952 (1 nymph,
reared DAC). Tennessee. Shelby County: 6 mi. N Memphis,
Microtus pinetorum, February 2, 1954 ( 1 nymph, reared DAC, idet.
by larval skin).

Genus Euschongastoides Loomis, New Status

Trombicula (Euschongastoides) Loomis, 1954, Univ. Kansas Sci. Bull., vol. 86,
p. 924 (type Trombicula (Euschongastoides) hoplai Loomis) (larva).

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates ab-
sent. (3) Body setae branched, not expanded, tips variable; with-
out compound branching; adjacent setae of similar lengths. (4)
Claws on tarsus I undi\dded distally. (5) Tarsus I without dorso-
apical stumplike process. (6) Cheliceral base not unusually elon-
gated; blade shearlike. (7) Palpal trochanter not elongated. (8)
Hypostome short, blunt, with about twenty short apical nude or
nearly nude setae. (9) Pharynx marked with a reticulate pattern.
(10) Tectum broad, hyaline, icith small serrations on margin, and
with one small tectal seta. ( 11 ) SensiJlary area wide; mean ratio
ASL/SB: 1.2-1.3. (12) Without branched body seta in sensillary
area. (13) Sensilla almost flagelliform but slightly thickened, with
branches.

Additional features (nymph). — (14) Body size medium (about
0.6 mm. long), of usual shape. (15) Sternum roughly rectangular,
closed behind. (16) Tarsus I but slightly elongated. (17) Articu-
lating area of cheliceral base with basis capituli posterior; base with
posterior tip elevated. (18) Palpus stout, small; distal portion of
tibia not unusually elongated; claw slightly curved, moderate in
size; two accessory claws; tarsus with five to seven branched setae
and two apical nude setae. ( 19 ) Hjqjostome with about ten ventral
branched setae. (20) Basis capituli short, broad. (21) Crista short,
narrow. (22) Sensillary area with basal plate of two well-separated
halves; bulla triangular, moderate in size; carina absent; sensillary
bases placed well lateral; ridges weak, apodeme apparently absent.
(23) About ten parascutal setae.

Remarks. — Euschongastoides was erected by Loomis as a sub-
genus of Trombicula for T, hoplai Loomis. At that time Loomis
noted that the affinities of T. hoplai appeared to be with certain
species of Euschongastia rather than with other species of Trom-

Nymphs of the Trombiculidae 227

hicula, despite the fact that the sensilla of T. hoplai ( as larva ) are
not expanded. The species of Euschongastia which are particularly
similar are E. finleyi Crossley, E. lacerta Brennan, and E. loomisi
Crossley and Lipovsky. Nymphs of E. loomisi were obtained.
It became evident that E. loomisi and T. hoplai were very similar
in the nymphal as well as in the larval stage, and that the nymphs
were abundantly distinct from those of other species of Trombicula
and Euschongastia.

This is certainly not the first instance of similar larval forms be-
ing separated in the two genera Trombicula and Euschongastia on
the basis of difference in sensilla (see Audy, 1954:133 concerning
confusion of larvae of T. munda and E. indica). In the present case,
however, the nymphs have been reared, and have been found to
show strong similarities to each other and appear to have aflSnities
with different genera from those in which the larvae were placed.

These two species present a taxonomic dilemma. To leave them
in the genera where they were originally placed would be in effect
to ignore evidence of the postlarval stages, not only by separating
two very similar nymphs but by placing them with groups which
are markedly different. To place the species on postlarval evidence
alone (perhaps in Pseudoschongastia, nymphs of which are quite
similar) would be to ignore evidence from the larval stage, in
particular the nature of the sensilla, which separates the two major
genera Trombicula and Euschongastia. After careful consideration
it appears that the flaw must lie in the rigid application of the
single larval character, flagelliform sensilla versus expanded sensilla.
Even in the larval stage the two species are similar except in the
sensilla. The solution which does the least violence to both sys-
tems of classification appears to be the recognition of full generic
status for Euschongastoides and the inclusion of E. loomisi in that
genus.

The genus Euschongastoides will be very difficult to recognize
in the larval stage until more species have been reared and a much
better understanding of its limits obtained. It is proposed that for
the time being no species be placed in this genus on the evidence
of either larval or postlarval stages alone.

The relationships of the genus Euschongastoides appear to be
with Pseudoschongastia and Walchia. Together, these three genera
are separable from other trombiculine genera by the following
diagnostic characters: (8) Hypostome short, blunt, with about
twenty short apical nude or nearly nude setae. Other genera have

228 The University Science Bulletin

the hypostome longer, usually pointed, with about eight apical
nude setae. (9) Pharynx marked with a reticulate pattern. In
other genera the pharynx is unmarked. (11) Mean ratio ASL/SB:
1.2-1.7. In other genera the mean ratio ASL/SB is 2.0 or greater
(except Gunthcrana and Doloisia, as described by other authors).
(17) Articulating area of cheliceral base with basis capituli pos-
terior, base with posterior tip elevated. In other genera the articulat-
ing area is on the ventral surface rather than across the posterior end.
The posterior tip of the cheliceral base is depressed. There are sev-
eral additional characters which indicate relationship between these
three genera. As previously discussed, these genera may form a
valid subfamily, but since larvae of Euschongastoides are at present
not separable from those of Trombicida and Euschongastia, recogni-
tion of such a subfamily seems impractical.

The genus Schoutedenichia, which I have not studied, appears
related to the Euschongastoides group of genera. Descriptions and
illustrations of nymphs of Schoutedenichia indicate that they share
the features listed above.

Euschongastoides is separable from Pseudoschongastia by the
nature of the sensilla (rather thin and quite flexible in Euschon-
gastoides, thicker and but slightly flexible in Pseudoschongastia)
and by the tectal margin (smooth in Pseudoschongastia, with small
teeth in Euschongastoides) . Walchia differs from Euschongastoides
in several characters, including the absence of a tectal seta and the
possession of a dorsoapical stumplike process on tarsus I in Walchia.

Nymphs of Euschongastia perameles (Womersley) and E. smithi
( Womersley ) have been described by Domrow ( 1955 ) . The de-
scriptions and illustrations indicate that these nymphs will probably
key out to Euschongastoides. Possibly, they would fall in Pseudo-
schongastia. At any rate, it is evident that these species belong
in the Euschdngasioides-Pseudoschongastia-Walchia group of gen-
era and not in Euschongastia. These species share with the Euschon-
gastoides group of genera the pertinent characteristics (8), (11),
and (17); two species possess precoxal plates and so violate char-
acter (2) for Euschongastoides and its relatives. Larvae of E.
perameles and E. smithi have been described by Womersley; scuta
of these larvae are similar to those of species of Euschongastoides.
I am uncertain as to whether these two species should be accom-
modated in Euschongastoides, Pseudoschongastia, or a new genus.

The nymph of Euschongastia indica (Hirst) as described by
Wharton (1946:159-161) suggests that this species may belong to

Nymphs of the Trombiculldae 229

Euschongastoides. Wharton's description is not sufficiently com-
plete to make placement positive.

Key to Species of Euschongastoides ( Nymphs )

1. Posterior body setae with tips ending in 2-4 attenuated branches;

tectal seta longer (about 12 jj,) with more branches Jwplai p. 229

2. Posterior body setae with tips rather abruptly pointed and bearing
6 or more very minute branches; tectal seta shorter (about 7 j;,)

with fewer branches loomisi p. 230

Euschongastoides hoplai (Loomis) New Combination

(Figs. 31, 87, 126, 127, 146, 179, 186, 219)

Trombicula {Euschongastoides) hoplai Loomis, 1954, Univ. Kansas Sci. Bull.,
vol. 36, pp. 294-296 (larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37,
pp. 1305-1306 (larva).

Description of nymph. — Body: About 0.6 mm. long, color white.
Setae rather short, with fine branches tending to project at right
angles from stems; tips of posterior setae ending in 2 to 4 short
attenuated branches; posterior setae about 35 \i long. Sternum
roughly rectangular in shape, rather distinctive, with 4 to 6 setae.
Measurements of tarsus I (means of 10 specimens); TL: 86.2 ±
3.095, TH: 50.9 ±: 0.924, TL/TH: 1.69 ± 0.0526.

Gnathosoma: Pharynx with reticulate markings covering about
half its width. Cheliceral base stout, distinctive in shape, punctate;
blade elongated but shearlike, apodeme of blade not constricted
and rather narrow; blade with about 9 rather small teeth commonl)^
on distal half. Cheliceral measurements (means of 12 specimens);
BL: 52.3 ± 1.534, BH: 25.6 ± 0.4345, CL: 24.8 ± 0.7054, BL/BH:
2.03 ± 0.0467, BL/CL: 2.10 ±: 0.0264. Palpus stout, small; acces-
sory claws thin; dorsal tibial setae large; tarsala about 6 (x long;
tarsus with 1 inner and 1 outer apical nude setae. Setal counts
(means of 7 specimens); femur: 6.93 ± 0.3996, genu: 8.00 ± 0.2182,
tibia: 5 (no variation), tarsus: range 6-7.

Scutum: Tectum with about 15-20 small serrations on margin;
seta short but small branches rather numerous. Crista very narrow.
Sensillary area with ridges weak but visible; no ridges medial to
sensillary bases; sensilla flexible; longer branches on distal part of
sensilla tend to run at right angles to stems, stems assume zigzag
appearances near apex. Scutal measurements (means of 10 speci-
mens): ASL: 52.2 ± 1.769, SB: 42.2 ± 1.872, TS: 12.2 ± 1.278,
SENS: about 75, ASL/SB: 1.26 ± 0.0372.

Remarks. — This species is apparently widely distributed in the

230 The University Science Bulletin

central and southwestern United States. Postlarvae probably in-
habit the soil surrounding the nests of their mammalian hosts.

Euschongastoides hoplai proved very difBcult to cultiu-e. The
inactive stages seem unusually susceptible to mold. The nymphs
move very sluggishly. Both Collembola eggs and crushed collem-
bolans were offered as food materials, but the nymphs did not eat.

An obvious question concerns the proper identification of the
nymphal stages of E. loomisi and E. hoplai. A field trip was made
to Barber County, Kansas, in 1955, with specific objective of obtain-
ing unquestionable evidence of the nymphal identification. In the
case of E. loomisi this was achieved, by the isolated rearing of a
single individual and recovery of the larval skin. Several individuals
of E. hoplai were found but none reached the nymphal stage. There
is, however, strong evidence associating the nymphal form with the
larvae of E. hoplai. This species was reared by me on two occa-
sions; in both instances larvae similar to those placed in culture were
identified as E. hoplai when the culture was started. Dead larvae
removed from the cultures were E. hoplai. No nymphs of this type
were reared in mixed cultures, nor were any found as contaminants
of other cultures. Also, the association of larva and nymph was ob-
served independently (and preceding my work) by Louis J. Lipov-
sky. Thus it appears most certain that the nymphs described as
E. hoplai are correctly identified.

Specimens examined. — Total 12 nymphs, as follows: Kansas.
Barber County: 1 mi. W, 4/2 mi. S Aetna, Cynomys ludovicianus, July
27, 1952 (7 nymphs, reared DAC); 10^2 mi. W Hardtner, Cynomys
ludovicianus, July 26, 1952 ( 1 nymph, reared DAC ) ; 4 mi. S Aetna,
Neotoma micropus, August 22, 1949 (4 nymphs, reared LJL).

Euschongastoides loomisi (Crossley and Lipovsky),

New Combination

(Figs. 8, 30, 223)

Euschongastia loomisi Crossley and Lipovsky, 1954, Proc. Ent. Soc. Washing-
ton, vol. 56, pp. 243-246 (larva); Loomis, 1956, Univ. Kansas Sci. Bull.,
vol. 37, pp. 1346-1347 (larva).

Description of nymph. — Resembles E. hoplai except as follows:
Body: About 0.6 mm. long. Setae with tips rather abruptly

pointed, with variable number (six or more) very tiny branches;

posterior setae about 40 [i long. Measurements of tarsus I (means

of 9 specimens); TL: 78.9 ± 1.006, TH: 46.4 ± 0.989, TL/TH: 1.70

rb 0.0236.
Gnathosoma: Cheliceral base with few puncta; blade with

apodeme shorter and broader; with about 9 teeth restricted to distal

Nymphs of the Trombiculidae 231

half of blade. Cheliceral measurements (means of 10 specimens);
BL: 51.5 ± 0.6709, BH: 24.2 ± 0.5541, CL: 24.2 ± 0.4899, BL/BH;
2.14 ± 0.0494, BL/CL: 2.14 ± 0.0394. Palpus with tarsala about
5 [X long. Setal counts; femur (8 specimens): 5.63 zt 0.2060, genu
(10 specimens): 7.3 ± 0.3267, tibia: range 4-6, tarsus: range 5-6.

Scutum: Tectal seta smaller, with fewer branches. Scutal meas-
urements (means of 8 specimens); ASL: 51.0 ± 1.662, SB: 46.1 ±
1.576, TS: 7.25 ± 0.4532, SENS: about 90, ASL/SB: 1.17 ± 0.0361.

Remarks. — Like E. hoplai, E. loomisi nymphs were inactive in
culture. They were not observed to feed on the Collembola eggs
and crushed collembolans offered to them.

Enschongast aides loomisi is known from the central and south-
western United States and from northern Mexico. Postlarvae prob-
ably inhabit the soil surrounding the nests and burrows of their
mammalian hosts.

Specimens examined. — Total 17 nymphs, as follows: Kansas.
Barber County: 4 mi. S Aetna, Neotoma micropus, July 11, 1955
(1 nymph, reared RBL, idet. by larval skin) and August 22, 1949
(13 nymphs, reared LJL), and Peromysciis manictdatiis, October
7, 1951 ( 1 nymph, reared LJL ) ; 3 mi. E, 5 mi. S Aetna, Neototna
micropus, July 25, 1952 ( 1 nymph, reared DAC ) ; 10^2 mi. W Hardt-
ner, Cynomys ludoviciamis, July 26, 1952 (1 nymph, reared DAC).

Genus Neoschongastia Ewing

Neoschdngastia Ewing, 1929, Manual of External Parasites, Springfield, 111.,
Thomas, p. 187 (type Schongastia americana Hirst) (larva); Wharton and
Hardcastle, 1946, Jour. Parasit., vol. 32, p. 288 (nymph); Wharton and
Fuller, 1952, Mem. Ent. Soc. Washington, no. 4, p. 84 (nymph); Womersley,
1952, Rec. South Australian Mus., vol. 10, p. 388 (nymph).

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates of
coxae I present but not fused in midline to form longitudinally di-
vided sternum. (3) Body setae with compound branching; not
expanded; long and short setae iniermixed. (4) Claws on tarsus
I undivided distally. (5) Tarsus I without dorsoapical stumplike
process. (6) Cheliceral base not unusually elongated; blade nar-
rowed but shcarlike. (7) Palpal trochanter not elongated. (8)
Hypostome projecting but blunt at tip; with about eight short
apical nude setae. (9) Pharynx indistinct. (10) Tectum large,
hyaline, with teeth; with one tectal seta. (11) Sensillary area
reniform; mean ratio ASL/SB: about 2.0-2.2. (12) Without
branched body seta in sensillary area. ( 13 ) Sensilla variable, with
branches.

232 The University Science Bulletin

Additional features (nymph). — (14) Body size medium (about
0.6 mm. long), strongly constricted, truncate posteriorly. (15)
Precoxal plates of coxae I touching in midline but not fused; sternum
open posterior to plates. (16) Tarsus I elongated. (17) Articulat-
ing area of cheliceral base with basis capituli ventral but short-
ened; base with posterior tip depressed. (18) Palpus unsually
stout, broad; usually elongated; claw curved, extremely elongated
in American species; one or two accessory claws; tarsus with about
nine nude or nearly nude setae and one apical nude seta. (19)
Hypostome with ten to twenty ventral branched setae. (20) Basis
capituli of usual shape. (21) Crista long but not narrowed, rod-
like. (22) Sensillary area with basal plate of two rather widely
separated halves; bulla much reduced in size; carina absent; sen-
sillary bases rather small, placed well forward; ridges present but
variable; apodeme much reduced. (23) About ten parascutal
setae.

Remarlis. — In addition to the nymphs described in the present
paper, others have been described. Wharton and Hardcastle (1946:
288) published descriptions of the nymphs of N. carveri Wharton
and Hardcastle and another species, either americana solomonis
Wharton and Hardcastle or monticola Wharton and Hardcastle.
The nymph of N. gallinarwn (Hatori) has been described by
Womersley (1952:390-391). The foregoing diagnosis is in agree-
ment with these published accounts, except as follows: The palpal
claws of other than American species are not extremely elongated.
No mention is made of the presence or absence of precoxal plates,
either by Wharton and Hardcastle or by Womersley. The single
nymph of N. carveri has two tectal setae and is probably aberrant
in this respect.

As larvae, species of Neoschongastia are recognized by the sub-
mergence of the scutum beneath the cuticular striae. The larvae
otherwise appear similar to many species now placed in Euschon-
gastia.

Although the larvae are placed by this single key character,
nymphs are abundantly distinct from those of other genera. It is
difficult to suggest to which genera Neoschongastia may be related.
The nymphal scutum is distinctive; it is somewhat suggestive of
Cheladonta but nymphs of these two genera seem specialized along
diflFerent lines and certainly do not share many other features. Of
the characters listed above, the elongated palpal claws are the most
ready means of identification for the American forms. Otherwise,

Nymphs of the Trombiculidae 233

the body setae are distinctive in being compoundly branched long
and short setae being intermixed.

Womersley (1952:389) gives an adequate key to the known
nymphs of Neoschongastia. American species are easily separated
from these by their very elongated palpal claws. The two Ameri-
can species are themselves separated in the following key.

Key to American Species of Neoschongastia ( Nymphs )

1. Cheliceral blade without teeth; palpal tibia with two accessory
claws; sensilla not expanded in midregions, flexible, with long
branches restricted to distal third brennani p. 234

2. Cheliceral blade with teeth; palpal tibia with one accessory claw;
sensilla somewhat expanded in midregion, with numerous spinehke
branches not restricted to distal third americana p. 233

Neoschongastia americana (Hirst)
(Figs. 40, 71, 177, 180, 193, 194, 201, 222, 224)

Schdngastia americana Hirst, 1921, Ann. and Mag. Nat. Hist., vol. 17, p. 37
( larva ) .

Neoschongastia americana, Ewing, 1929, Manual of External Parasites, Spring-
field, 111., Thomas, p. 187 (larva); Loomis, 1956, Univ. Kansas Sci. Bull.,
vol. 37, pp. 1354-1355 (larva).

Description of nymph. — Body: About 0.6 mm. long, color red.
Setae with thick stems and short thick branches which in turn di-
vide into 2-4 fine branches; tips of posterior setae with 2-3 shorter
compound branches; longer body setae about four times as long as
shorter ones; long posterior setae about 70 [i. long. Sternum with
rounded precoxal plates of coxae I meeting at midline, each plate
bearing 2-4 setae; sternal area open behind. Legs appear short-
ened. Measurements of tarsus I (means of 10 specimens); TL:
95.4 ± 0.9214, TH: 49.0 ± 0.5164, TL/TH: 1.94 ± 0.0305.

Gnathosoma: Hypostome with about 8 short apical nude setae.
Cheliceral base normal, punctate; blade narrowed, elongated but
shearlike, with about 10 teeth. Cheliceral measurements (means
of 10 specimens); BL: 71.6 ± 1.845, BH: 28.7 ± 0.7000, CL: 30.6
± 0.5927, BL/BH: 2.51 ± 0.0781, BL/CL: 2.36± 0.0851. Palpus
with claw very elongated, dorsal tibial seta heavy; lower accessory
claw absent, apparently replaced by branched body seta; upper ac-
cessory claw normal; tarsala about 8 [i long; most tarsal setae nude,
rather thick; tarsus with apical nude seta elongated, on outer sur-
face. Setal counts; femur (8 specimens): 3.38 ±: 0.1813, genu (9
specimens) : 7.83 ± 0.2041, tibia: range 3-5, tarsus, range 7-9.

Scutum: Tectum nearly square. Sensillary area with basal plate
rather distinctive; ridges weak but obvious, lending rcniform ap-

234 The University Science Bulletin

pearance to area; bulla small and square; sensilla thick, somewhat
expanded in midregions, with numerous spinelike branches. Scutal
measurements (means of 9 specimens); ASL: 62.3 d= 0.6872, SB:
28.7 ± 2.0178, TS: 44.1 dz 1.2427, SENS: about 90, ASL/SB: 2.17
± 0.0532.

Remarks. — Neoschongastia americana appears to be a widespread
species in the southeastern United States and in other areas. Most
of the hosts of this species are birds; however, Loomis (1956:1355)
shows that cottontails are a very important host animal. Loomis
also presents evidence to show that the postlarvae probably inhabit
soils which are well drained, warm to hot and dry.

Larvae of N. americana and N. brennani are quite similar, but
nymphs of these species diflFer in several characters, as indicated in
the key. DifiFerences between the two species were first observed
in the nymphal stage by L. J. Lipovsky. Wharton and Hardcastle
(1946:313) describe a nymph which is either that of N. americana
solomonis or of N. monticola. There are numerous differences be-
tween their description and the nymph of N. americana. Probably
the nymph pertains to IV. monticola; if it is indeed the nymph of
N. a. solomonis, then solomonis should be considered a distinct
species.

Neoschongastia americana proved very diflBcult to culture. Mod-
erate success was achieved in culture tubes which were allowed to
remain much drier than usual. The nymphs ate active stages of
collembolans, but ignored their eggs.

Specimens examined. — Total 38 nymphs, as follows: Kansas.
Douglas County: Lawrence, Sylvilagus floridanus, July 11, 1949
(10 nymphs, reared LJL), and Colinus virginianus, September 15,
1952 ( 10 nymphs, reared RBL-DAC ) ; 4 mi. S Lawrence, Sylvilagus
■floridanus, July 25, 1948 (2 nymphs, reared LJL); 5 mi. S Lawrence,
Sylvilagus floridanus, July 14, 1948 (16 nymphs, reared LJL).

Neoschongastia brennani Crossley and Loomis

(Figs. 11, 41, 72, 100, 101, 149)

Neoschongastia brennani Crossley and Loomis, 1955, Ent. News, vol. 66, pp.
114-117 (larva).

Description of nymph. — Resembles N. americana except as fol-
lows:

Body: About 0.6 mm. long. Setae with somewhat longer
branches; posterior setae about 70 [j. long. Precoxal plates with each
plate bearing about 8 setae. Measurements of tarsus I (means of

Nymphs of the Trombiculidae 235

2 specimens); TL: 91.5 ± 3.50, TH: 44.5 ± 0.50, TL TH: 2.05
± 0.050.

Gnathosoma: Hypostome with about 8 very short apical nude
setae, Chehceral blade without teeth. Cheliceral measurments
(means of 2 specimens); BL: 94.5 ± 0.50, BH: 34.5 ±: 2.50, CL:
36.5 ± 1.50, BL/BH: 2.8 ± 0.250, BL/CL: 2.6 ± 0.140. Palpus
with lower accessory claw present but small; tarsala about 7 [x long.
Setal counts (means of 2 specimens); femur: 3.5 ± 0.000, genu:
10.25 ± 1.2502, tibia: 3 (all on outer surface), tarsus: range 8-9.

Scutum: Sensilla thick, not expanded in midregions. flexible;
with long branches restricted to distal thirds. Scutal measurements
(means of 2 specimens); ASL: 66.0 ± 8.000, SB: 34.0 ± 3.00, TS:
38.0 ± 0.00, SENS: about 115, ASL/SB: 1.95 ± 0.500.

Remarks. — Loomis (1956:1359) notes that this species is known
only from birds, particularly the woodpecker, Melonerpes cnjthro-
cephalus, and suggests that postlarvae may inhabit standing or
fallen dead trees.

Specimens examined. — Total 2 nymphs, as follows: Kansas.
Barber County: 10/2 mi. W Hardtner, Melanerpes enjtlirocephalus,
July 26, 1952 (2 nymphs, reared DAG).

Genus Psetidoschongastia Lipovsky

Pseudoschdngastia Lipovsky, 1951, Jour. Kansas Ent. Soc, vol. 24, p. 95 (type
Pseudoschongastia hungerfordi Lipovsky) (larva).

Diagnosis (mjmph). — (1) Eyes absent. (2) Precoxal plates ab-
sent. (3) Body setae branched, not expanded, without compound
branching; adjacent setae of similar lengths. (4) Claws on tarsus
I undivided distally. (5) Tarsus I without dorsoapical stumplike
process. (6) Cheliceral base not unusually elongated; blade shear-
like. (7) Palpal trochanter not elongated. (8) Hypostome short,
blunt; with about twenty short apical nude or nearly nude setae.
(9) Pharynx marked with a reticulate pattern. (10) Tectum broad,
hyaline, with smooth margin; with one small tectal seta. (11)
Sensillary area wide, mean ratio ASL/SB: about 1.7. (12) With-
out branched body seta in sensillary area. (13) Sensilla straight,
rather thick to subclavate, with branches.

Additional features (nymph). — (14) Body medium in size (about
0.6 mm. long), elongated. (15) Sternum broadly pentagonal,
closed behind. (16) Tarsus I elongated. (17) Articulating area
of cheliceral base with basis capituH posterior; base with posterior
tip depressed. (18) Palpus stout, small; distal portion of tibia not

236 The University Science Bulletin

unusually elongated; claw slightly curved, moderate in size; two
accessory claws; tarsus with four to seven branched setae and two
apical nude setae. (19) Hypostome with about ten ventral
branched setae. (20) Basis capituli short, broad. (21) Crista
short, narrow. (22) Sensillary area with basal plate of two well-
separated halves; bulla rather large; carina absent; sensillary bases
placed well laterally; anterior, lateral, and posterior ridges weak;
ridges medial to sensillary bases absent; apodeme apparently ab-
sent. (23) About ten parascutal setae.

Remarks. — Fseudoschongastia is separable from, but closely re-
lated to, Eiischongastoides. Also, its relationship to Walchia and to
Schotdedenichia is evidenced by several characters. Audy (1954:
155) suggests that Psetidoschongastia is really related to Euschon-
gastia, an opinion which I do not share. However, Audy may be
considering as typical Eiischongastia certain species which would
better be accommodated in Eiischongastoides.

Larvae of Fseudoschongastia are recognized by the posterolateral
scutal setae being displaced from the scutum and the legs II and
III having their femurs undivided. Otherwise, they also show
similarities to larvae of Euschongastoides.

In culture, both species of Fseudoschongastia eat active stages
of collembolans rather than their eggs.

Key to Species of Pseudoschongastia (Nymphs)

1. Reticulate markings cover about half the width of the pharynx;
chehceral blade with about 6 large conspicuous teeth; sensilla
straight, never expanded farneri p. 237

1'. Reticulate markings restricted to midline of pharynx; cheliceral
blade with about 10 small teeth; sensilla straight, usually some-
what expanded but occasionally not hungerfordi p. 236

Fseudoschongastia hungerfordi Lipovsky

(Figs. 32, 91, 144, 218)

Fseudoschongastia hungerfordi Lipovsky, 1951, Jour. Kansas Ent. Soc, vol.
24, pp. 95-99 (larva).

Description of nymph. — Body: About 0.6 mm. long, color white.
Setae short, with long branches; tips of posterior setae composed
of 2-3 rather long branches; posterior setae about 20 [a long. Ster-
num closed behind, roughly pentagonal, with about 8 setae. Meas-
urements of tarsus I ( means of 9 specimens ) ; TL: 88.1 ± 2.300, TH:
52.1 ± 1.263, TL/TH: 1.70 ± 0.0373.

Gnathosoma: Reticulate markings on pharynx restricted to mid-
line. Cheliceral base stout, puncta apparently absent; blade shear-
like, with about 10 small teeth covering most of dorsal margin;

Nymphs of the Trombiculidae 237

occasional specimens with fewer ( about 8 ) larger teeth. Cheliceral
measurements (means of 8 specimens); BL: 44.5 =h 2.535, BH:
23.3 ± 0.974, CL: 26.6 ± 0.681, BL/BH: 2.01 ± 0.106, BL CL:
1.76 rf= 0.0906. Palpus stout, small, accessory claws thin; tarsala
about 7 [X long; tarsus with one inner and one outer apical nude
seta. Setal counts (means of 8 specimens); femur: 3.00 ± 0.125,
genu: 4.10 ± 0.164, tibia: range 2-3, tarsus: range 4-6.

Scutum: Tectal seta small and with few branches (about 4).
Sensillary area with bulla triangular; sensilla stiff, usually somewhat
expanded and nearly subclavate, rarely unexpanded, vith short
branches on basal thirds, long straight thick-based branches on
midregions, and shorter but similar branches on apical thirds.
Scutal measurements (means of 10 specimens); ASL: 55.3 ±
0.9781, SB: 32.3 ± 0.6047, TS: 6.75 ± 0.250, SENS: about 80,
ASL/SB: 1.69 ± 0.0378.

Remarks. — The characters in the key separate most specimens
of P. himgerfordi from P. farneri. Several poorly mounted speci-
mens could not be placed satisfactorily in one species or the other.

Psetidoschongastia himgerfordi was readily cultured, and a num-
ber of adults were so obtained.

Specimens examined. — Total 149 nymphs, as follows: Kansas.
Barber County: 3 mi. S Aetna, Neotoma microptts, July 25, 1952
(5 nymphs, reared DAC); 4 mi. S Aetna, Neotoma micropiis, July
11, 1955 (7 nymphs, reared DAC, idet. by larval skins), and
August 22, 1949 (46 nymphs, reared LJL); 5 mi. S, 3 mi. E Aetna,
Neotoma micropiis, July 25, 1952 (2 nymphs, reared DAC). Doug-
las County: 5 mi. S Lawrence, Sylvilagus floridanus, July 14, 1948
(10 nymphs, reared LJL). Greenwood County: VA mi. E Hamil-
ton, Neotoma jloridana, August 14, 1949 (46 nymphs, reared LJL).
Rawlins County: 9 mi. W Atwood, Reithrodontomys megalotis,
August 10, 1949 (2 nymphs, reared LJL). Texas. Bexar County:
Camp Bullis, Sylvilagus auduboni, April 25, 1954 (31 nymphs,
reared RBL-DAC).

Pseudoschongostia farneri Lipovsky

(Figs. 33, 90, 124, 125, 145)

Psetidoschongastia farneri Lipovskv, 1951, Jour. Kansas Ent. Soc, vol. 24, pp.
101-102 (larva).

Description of nymph. — Resembles P. hungerfordi except as fol-
lows :

Body: Measurements of tarsus I (means of 9 specimens); TL:
90.0 ± 2.06, TH: 48.9 ± 1.07, TL TH: 1.88 ± 0.0325.

238 The University Science Bulletin

Gnathosoma: Reticulate markings covering about half the width
of the pharynx. Cheliceral blade with about 6 larger conspicuous
teeth; sometimes with more (about 8) teeth. Cheliceral measure-
ments (means of 9 specimens); BL: 55.8 =h 0.7412, BH: 29.4 zt
0.7287, CL: 29.9 ± 0.5880, BL/BH: 1.91 ± 0.2609, BL/CL: 1.88 ±
0.2146. Setal counts of palpus (means of 9 specimens); femur:
2.89 ± 0.139, genu: 6.17 ± 0.238, tibia: range 3-4, tarsus: range
4-7.

Scutum : Sensilla like those of P. hungerfordi but never expanded.
Scutal measurements (means of 9 specimens); ASL: 54.3 ± 1.10,
SB: 32.7 ± 1.146, TS: 6.25 ± 0.500, SENS: about 70, ASL/SB:
1.69 ± 0.0696.

Remarks. — Unlike P. hungerfordi, P. farneri was very difficult
to maintain in culture. This suggests that P. farneri has different
requirements, which were not met in the cultures. Loomis (1956:
1350) suggests, however, that these two species are probably often
in direct competition as postlarvae, since they frequently occur
on the same host and have about the same range of host species.

Specimens examined. — Total 13 nymphs, as follows: Kansas.
Barber County: 3 mi. S Aetna, Neotoma micropus, July 25, 1952 (2
nymphs, reared DAC ) ; 3/2 mi. S, 1 mi. W Aetna, Neotoma micropus,
April 11, 1949 (1 nymph, reared LJL); 4 mi. S Aetna, Neotoma
micropus, July 11, 1955 (4 nymphs, reared DAC, idet. by larval
skins). Lyon County: 2 mi. S Chalk, Perognathus hispidus. May
31, 1950 (1 nymph, reared LJL). Russell County: 9 mi. S Russell,
Peromyscus maniculatus, April 26, 1952 (4 nymphs, reared DAC);
5 mi. N, 2 mi. E Graham, Peromyscus maniculatus, April 27, 1952
( 1 nymph, reared DAC ) .

Genus Speleocola Lipovsky

Speleocola Lipovsky, 1952, Jour. Kansas Ent. Soc, vol. 25, p. 134 (type Speleo-
cola tadaridae Lipovsky) (larva); Audy, 1954, Stud. Inst. Med. Res. Fed.
Malaya, no. 26, p. 147 (larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol.
37, p. 1326 (larva).

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates of
coxa I absent. (3) Body setae short, not expanded; with compound
branching; adjacent setae of similar lengths. (4) Claws on tarsus I
undivided distally. (5) Tarsus I without dorsoapical stumplike
process. (6) Cheliceral base unusually elongated (ratio BL/BH:
about 3.5); blade shearlike. (7) Palpal trochanter not elongated.
(8) Hypostome pointed, projecting, with about eight apical nude
setae. (9) Pharynx indistinct. (10) Tectum small, hyaline, margin

Nymphs of the Trombiculidae 239

with teeth; with one tectal seta. (11) Sensillary area triangular;
ratio ASL/SB: about 2.5. (12) Without branched body seta in
sensillary area. (13) Sensilla subclavate; branches short with heavy
bases.

Additional features (nijmph). — (14) Body small (about 0.4 mm.
long). (15) Sternum not closed behind, somewhat elongated.
(16) Tarsus I elongated. (17) Articulating area of cheliceral base
with basis capituli ventral; posterior tip of cheliceral base depressed.
(18) Palpus somewhat elongated, size small; distal portion of tibia
not unusually elongated; claw slightly curved, somewhat elongated;
two accessory claws; tarsus with about nine nude or nearly nude
setae and three apical nude setae. (19) Hypostome with about
ten ventral branched setae. (20) Basis capituli elongated. (21)
Crista a narrow rod. (22) Sensillary area with basal plate of two
halves; bulla reduced in size; carina present; sensillary bases placed
well lateral; anterior and lateral ridges strong but posterior ridge
weak; ridges medial to sensillary bases bearing small carinae; apo-
deme normal. ( 23 ) About eight parascutal setae.

Remarks. — Speleocola is monotypic for S. tadaridae. As a larva
this species has a peculiar scutum and particularly peculiar sensilla.
The sensilla are apparently unexpanded but the branches are broad
based and leaflike, possibly producing the effect of an expanded
sensillum.

The nymphs of S. tadaridae are quite similar to those of Trombic-
ula species. The outstanding differences are the compound branch-
ing of the body setae and the carinate ridges medial to the sensillary
bases. Whether or not these differences are sufficient to merit the
generic distinction is a debatable question.

Speleocola tadaridae Lipovsky

(Figs. 39, 86, 128,129, 173, 233)

Speleocola tadaridae Lipovsky, 1952, Jour. Kansas Ent. Soc, vol. 25, pp. 134-
137 (larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1326-1327
( larva ) .

Description of nymph. — Body: About 0.4 mm. long, color not
known. Setae short, with few branches further dissected into 2-4
compound branches; posterior setae about 12 \>. long. Sternum with
about 4 setae. Measurements of tarsus I ( means of 10 specimens ) ;
TL: 63.8 ± 0.442, TH: 28.4 ± 0.371, TL/TH: 2.25 ± 0.0428.

Gnathosoma: Cheliceral base slightly curved, elongated, area of
articulation covering about two-thirds of ventral margin, base
punctate; blade shearlike, rather elongated, with about 16 tiny

240 The University Science Bulletin

teeth. Cheliceral measurements (means of 10 specimens) BL:
91.7 ± 0.8172, BH: 26.6 ± 0.6531, CL: 37.4 ± 0.5413, BL/BH:
3.46 ± 0.0670, BL/CL: 2.45 ± 0.0268. Palpus small, with reduced
setation, and setae with reduced branching; dorsal setae of genu
and tibia nude or nearly so; tibial seta at inner tarsal articulation
nude; claw and accessory claws slender; tarsala about 4 \t. long;
tarsus with 2 inner, 1 outer apical nude setae. Setal counts ( means
of 10 specimens); femur: 3.45 ± 0.279, genu: 3.25 dz 0.1535, tibia:
range 4-5, tarsus: range 7-9.

Scutum: Tectum rather rounded; marginal teeth conspicuous;
seta near end of crista. Sensillary area with basal plate rather
small; apodeme present, normal, hard to see; bulla triangular and
small; carina thin; "saddle" portion of area large, punctate; sensilla
expanded, with numerous short spinelike branches. Scutal meas-
urements (means of 10 specimens); ASL: 54.9 ± 0.8175, SB: 22.4
±0.4521, TS: 14.4 ± 0.1633, SENS: about 55, ASL/SB: 2.45
± 0.0342.

Remarks. — This species was reared on a single occasion by L. J.
Lipovsky. The nymphs did not eat, and none reached the adult
stage.

Specimens examined. — Total 23 nymphs, as follows: Oklahoma.
Woods County: 6 mi. S, 2 mi. W Aetna, Kans., Tadarida mexicana,
August 24, 1949 ( 23 nymphs, reared LJL ) .

Genus Trombictda Berlese

Trombicula Berlese, 1905, Redia, vol. 2, p. 155 (type Trombicula minor Ber-
lese) (adult); Womersley and Heaslip, 1943, Trans. Roy. Soc. South Aus-
tralia, vol. 67, p. 73 (part) (adult); Ewing, 1949, Jour. Washington Acad.
Sci., vol. 39, p. 235 (adult); Wharton et al, 1951, Jour. Parasit., vol. 37, p.
30 (adult); Womersley, 1952, Rec. South AustraUan Mus., vol. 10, p. 326
(adult); Audy, 1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 137
( adult ) .

Diagnosis (nymph). — (1) Eyes absent or 1/1 and adjacent to
sensillary area. (2) Precoxal plates of coxae I absent or fused in
midline to form longitudinally divided sternum. (3) Body setae
variable, branched, without compound branching; adjacent setae of
similar lengths. (4) Claws on tarsus I undivided distally. (5)
Tarsus I without dorsoapical stumplike process. (6) Cheliceral
base rarely unusually elongated (mean ratio BL/BH: up to 3.6);
blade shearlike. (7) Palpal trochanter not elongated. (8) Hypo-
stome pointed, projecting; with about eight apical nude setae. (9)
Pharynx indistinct. ( 10 ) Tectum square to triangular, hyaline, mar-
gin with teeth; with one tectal seta. (11) Sensillary area triangular;

Nymphs of the Trombiculidae 241

mean ratio ASL/SB: about 2.0-2.7. (12) Without branched body
seta in sensillary area. (13) Sensilla variable, flagelliform to sub-
clavate, with at least basal branches.

Additional features (nymph). — (14) Body small to large (about
0.5-1.0 mm. long), sometimes elongated. (15) Sternum nearly
rectangular when longitudinally divided; otherwise open or closed
behind and elongated to roughly pentagonal. (16) Tarsus I
elongated. (17) Articulating area of cheliceral base with basis
capituli ventral; posterior tip of cheliceral base depressed. (18)
Palpus stout to somewhat elongated; distal portion of tibia not
unusually elongated; two accessory claws; tarsus with about nine
branched setae and three to six apical nude setae. ( 19 ) Hypostome
with about ten to twenty ventral branched setae. (20) Shape of
basis capituli variable. (21) Crista a narrow to very narrow rod.
( 22 ) Sensillary area with basal plate of two halves, variable; bulla
usually large, sometimes reduced; carina present or absent; sen-
sillary bases placed well lateral; anterior, lateral, and posterior
ridges present but variable; often additional ridges present; apo-
deme present, well developed to reduced. (23) About eight to
thirty parascutal setae.

Remarks. — In addition to the species described in the present
paper, postlarvae of about sixty species of Trombicula have been
previously described; these are discussed under the appropriate
subgenera. Most of the descriptions are inadequate but to my
knowledge none will be excluded from the genus by my diagnosis.
It should be noted, however, that the foregoing diagnosis is very
broad. Many previously described forms are known only as adults;
their placement is questionable. I feel that they should be left
in Trombictda until they can be definitely placed.

The type of the genus, Troinbicula minor Berlese, is known from
the original description of Berlese and a redescription of several
years later; Willmann in 1941 redescribed the types which were
subsequently destroyed by war. The type locality is a rather in-
accessible bat cave in Java; it has not been possible to collect
topotypical material. This mite has been the subject of much
discussion which I will not attempt to summarize; this has been
done by Womersley (1952:328-330) and by Audy (1954:138-140).
I shall attempt to list what definite information we do have con-
cerning T. minor. (1) The types were adults (as shown by Will-
mann) and almost certainly trombiculids. (2) The adults were
small, even for trombiculids. (3) The presence of these mites

242 The University Science Bulletin

in bat caves strongly suggests that the larvae are parasites of bats,
but this is not certain. (4) In the way of descriptive material we
have the brief descriptions of Berlese and of Willmann, and illus-
trations of scuta and palpi by these authors. (5) It seems safe to
assume that if T. minor possesses any strikingly unsual features these
would have been detected by either Berlese or Willmann.

The descriptions are quite brief. The illustrations of palpi are
difficult to evaluate; the palpi were drawn in a half-turned position
and it seems to me that they could represent almost any trombiculid.
The illustrations of the scutum by Berlese are poor; the illustra-
tion by Willmann is better but is obviously diagrammatic. Authors
have been unable to agree as to whether Willmann's drawing of
the sensillary area showed enlarged sensillary bases or eyes adja-
cent to the scutum (Willmann himself was uncertain on this point).
While Willmann's illustration appears to me to show a Trombicula
or Eitschdngastia sensillary area, I believe that with a slight stretch
of the imagination it could be fitted to several other genera.

From these considerations I draw the following conclusions.

(1) It is almost a certainty that T. minor is a small trombiculid.

(2) It is highly probable that the larvae are parasites of bats. (3)
It is possible that T. minor is a member of the genus Trombicula
as currently understood, but it could easily be considered con-
generic with species now placed in difi^erent genera. ( 4 ) It will be
impossible to recognize T. minor on the basis of the existing descrip-
tions.

As type of the genus Trombicula, T. minor is bound to continue
to cause taxonomic indecision and confusion. One acceptable solu-
tion would be to restrict Trombicula to T. minor and remove all
other species to a different genus (or genera). Concerning this
course, Wharton and Fuller (1951:42) write, "Such a course would
be entirely unjustified, since, despite the plethora of generic names
that have been proposed, Trombicula as understood here is no
broader in its content than are such genera as Neoschotiga.stia and
Euschongastia. In handling nomenclatorial problems of this kind
stability should be maintained if it is at all possible to do so and
still remain in agreement with the known biological facts." This
conservative course has so far been more productive than the alter-
native would be. However, the genus Trombicula has become
unwieldly, not through the number of species it contains but through
their diversity. At present the revision of this genus would con-
stitute a very difficult task, and as more species which key out to
Trombicula are added, the difficulties will increase. It seems to

Nymphs of the Trombiculidae 243

me that the best course would be to appeal to the International
Commission of Zoological Nomenclature to use its plenary powers
to stabilize the genus Trombicula, by replacement of the tj-'pe T.
minor with some well-known species. I would suggest that the
species selected be a bat parasite in the Asiatic-Pacific area.

The genus Trombicula has been divided into several subgenera;
there are more or less distinct groups of species which have been
left in Tro7nbictila primarily because they are possibly congeneric
with T. minor. Included in this paper are representatives of
Eutrombicula, Neotrombiciila, and Leptotrombidiwn, plus species
which fall into no well defined subgenus and are therefore cus-
tomarily placed in the subgenus Trombicula. These subgenera will
be discussed further under their headings.

As a genus, Trombicula as understood here is close to the genera
Speleocola, Blankaartia, and Euschongastia. Speleocola (mono-
typic) may well be an aberrant Trombicula. Blankaartia has prim-
itive characteristics but is nevertheless similar to some species of
Trombicula. Euschongastia is separable from Trombicula in both
larval and postlarval stages on single characters; Trombicula larvae
have unexpanded sensilla and nymphs have teeth on the tectal
margin, while Euschongastia larvae have expanded sensilla and
have a smooth tectal margin. Otherwise the genera have some
species which appear similar and it may become increasingly diffi-
cult to separate these two genera as they are currently understood.

Key to Species of Trombicula (NY^^'Hs)

1. Eyes present adjacent to sensillary area; posterior scutal ridge
with two strong elevations medial to sensillary bases

(subgenus EutromhicuJa) 2
1'. Eyes absent; posterior scutal ridge without elevations or with only
very weak elevations 5

2. Posterior body setae with tips rounded, slightly expanded, bear-
ing many short branches .- belkini p. 248

2' Posterior body setae tapering gradually, ending in 2-3 short
branches 3

3. Smaller; mean ASL about 90 jx, individual ASL less than 100 ^

alfreddugesi p. 245
3'. Larger; mean ASL about 115 jj. or more, individual ASL more
than 100 [J. 4

4. Posterior body setae mostly ending in two tiny branches, but

some ending in three lipovskyana p. 249

4'. Posterior body setae mostly ending in three tiny branches, but

some ending in two splendens p. 250

5. Precoxal plates of coxae I present and fused medially to form a
longitudinally divided sternum

(Subgenus Leptotrombidium) mijotis p. 257

244 The University Science Bulletin

5'. Precoxal plates of coxae I absent 6

6. Palpal tarsus with three apical nude setae (subgenus Trombicula,
part) 7

6'. Palpal tarsus with more than three apical nude setae 10

7. Cheliceral base unusually elongated (mean ratio BL/BH: 3.6)

ornata p. 267
7'. Cheliceral base not unusually elongated (mean ratio BL/BH:
less than 3.0) 8

8. Posterior body setae shorter, about 15 jj, long merrihewi p. 264

8'. Posterior body setae longer, about 20-25 ^ 9

9. Smaller species; ASL about 59 |j, crossleyi p. 258

9'. Larger species; ASL about 66 [j, trisetica p. 268

10. Palpal tarsus with four apical nude setae (T. ) montanensis p. 265

10'. Palpal tarsus with five or six apical nude setae 11

11. Sensilla nude except for short basal branches; rarely furcate at

tips ( ^. ) whartoni p. 255

11'. Sensilla with at least several distal branches in addition to small
basal branches 12

12. Sensilla flagelliform, very flexible, scarcely thicker than branches;
branches long (subgenus Trombicula, part) 13

12' Sensilla flexible but thickened, much thicker than branches;
branches usually shorter 14

13. Cheliceral base unusually elongated (BL/BH: 3.0) . . kansasensis p. 262
13'. Chehceral base not unusually elongated (mean ratio BL/BH:

2.6) gurneyi p. 261

14. Apical third of posterior body seta with many very short branches;
longer branches mostly restricted to basal portion

(subgenus Trombicula, part) 15
14'. Posterior body seta normal, not so modified

(subgenus Neotrombicida, part) 16

15. Occasional long branches on apical third of posterior body

seta kardosi p. 263

15'. Longer branches entirely restricted to basal portion of posterior

body seta fitchi p. 260

16. Smaller species; mean ASL about 65 [x sylvilagi p. 254

16'. Larger species; mean ASL about 84.97 y^ 17

17. Posterior body setae longer ( about 70 ^) autumnalis p. 252

17'. Posterior body setae shorter ( about 45 ^) lipovskyi p. 253

Subgenus Eutrombicula Ewing

Eutronibicula Ewing, 1938, Jour. Washington Acad. Sci., vol. 28, p. 293 (type

Microtrombidium alfreddugesi Oudemans) (larva).
Trombicula (Eutrombicula), Jenkins, 1949, Ann. Ent. Soc. America, vol. 42,

pp. 289-317 (postlarvae).

Diagnosis (nymph). — (1) Eyes 1/1, adjacent to sensillary area.
(10) Tectum square, with numerous small teeth on margin. (22)
Sensillary area with posterior scutal ridge with two strong eleva-
tions medial to sensillary bases. (23) About twenty to thirty para-
scutal setae.

Nymphs of the Trombiculidae 245

Remarks. — This subgenus could easily be raised to generic rank.
I have not done so, for two reasons: One, despite differences.
Eutrombicula species are very close to some other species of
Trombicula (for example, T. gurneyi); and two, species in the
Asiatic-Pacific area require re-examination (see Audy 1954:146).
Of the subgeneric characters the presence of eyes adjacent to the
sensillary area is probably the most important. However, it should
be pointed out that the eyes of T. belkini were seen on but one
specimen. Also, T. frittsi Wharton appears to have eyes in this
position (nymph described by Womersley, 1952, p. 330) but other-
wise does not seem to be a Eutro7nbicuIa. Nevertheless, I am in-
clined to look upon any nymph with eyes in this position as very
probably being a member of this subgenus. In culture, nymphs
readily ate Collembola eggs. Adults were obtained of all four
species cultured.

Other postlarvae of Eutrombicula have been described. The
species T. coarctafa Berlese, T. manriquei Ewing, and T. mediocris
Berlese are known as adults only. Nymphs of T. hirsti Sambon and
T. sarcina Womersley have been described. Both nyinphs and
adults of T. batatas (Linnaeus), T. samboni Womersley, and T.
wichmanni (Oudemans) have been described. I have not been
able to construct a key to handle these forms; the included key
contains only the species I have actually examined. Womersley
(1952:325) considers Willmann's drawing of T. minor to exhibit
eyes; therefore he considers Eutrombicula a synonym of Trombicula
(Trombicula). While this is possible, I prefer to follow the inter-
pretation of Ewing and others that the illustration shows enlarged
sensillary bases rather than eyes.

Trombicula alfrcddugesi (Oudemans)
(Figs. 21, 74, 110, 111, 168, 191, 226, 227)

Microthrcymhidium alfreddii^esi (Oudemans), 1910, Ent. Berichten, vol. .3, p.

84 (larva).
Eutrombicula alfreddugtsi, Ewing. 1938, Jour. Washington Acad. Sci., vol. 28,

p. 294 (larva).
Trorrthicula (Eutrombicula) alfrcddugesi, Jenkins, 1949, Ann. Ent. Soc. America,

vol. 42, p. 806 (adult); Wolfenbarger, 1953, Ann. Ent. Soc. America, vol.

45, pp. 652-654 (postlarvae).
Trombicula (Trombicula) alfreddugesi, Womersley, 1952, Rec. South Australian

Mus., vol. 10, p. 333.
Trombicula irritans, Ewing, 1929, Manual of External Parasites, p. 23 (adult).
Trombicula cinnabaris Ewing, 1921, Ann. Ent. Soc. America, vol. 13, p. 387

(adult).
Eutrombicula vanommereni, Michener, 1946, Ann. Ent. Soc. America, vol. 39,

p. 414 (postlarvae); Womersley, 1952, Rec. South Australian Mus., vol. 10,

p. 332.

Description of mjmph. — Body: About 0.9 mm. long engorged,
rather rounded posteriorly, color orange to red. Eyes rounded,

246 The University Science Bulletin

red, closely adjacent to sensillary area, about 17 [t. in diameter.
Setae long, stems thick but tapered, with many fine branches; tips
of posterior body setae with 2, rarely 3, short branches; posterior
setae about 60 [j. long. Sternum closed behind, roughly pentagonal
in shape, with about 8 setae. Measurements of tarsus I (means of
20 specimens); TL: 120.0 ± 2.028, TH: 65.1 ± 0.8032, TL TH:
1.85 ± 0.0246.

Gnathosoma: Chelicerae of usual facies; base punctate; blade
with about 20 teeth of various sizes. Cheliceral measurements
(means of 16 specimens); BL: 107.2 ± 2.049, BH: 46.6 ± 0.8803,
CL: 61.9 ± 1.3735, BL/BH: 2.28 ± 0.0393, BL/CL: 1.68 ± 0.0542.
Palpus of usual shape, except femur unusually stout; tarsala about
12 [t. long; tarsus with 3 inner, 2 outer apical nude setae. Setal
counts (means of 9 specimens); femur: 5:61 ± 0.2324, genu:
10.56 ± 0.3783, tibia: range 4-5, tarsus: range 9-10.

Scutum: Tectum about square, margin with about 20 small
teeth. Sensillary area with basal plate of two widely separated
bands; apodeme conspicuous; ridges strong; ridges medial to sen-
sillary bases elevated and joining posterior margin to give posterior
portion of sensillary area a 'lailobed" appearance; bulla triangular,
punctate, rather prominent; carina usually doubled, rather thick,
prominent; sensilla flagelliform, elongate, with short basal branches
and long attenuate distal branches; distal portions unipectinate.
Parascutal setae numerous, about 20-30. Scutal measurements
(means of 21 specimens); ASL: 88.1 ± 1.273, SB: 34.3 ± 0.4660,
TS: 34.5 ± 0.5986, SENS: about 130, ASL/SB: 2.58 ± 0.0318.

Remarks. — The synonomy given is far from complete, but the
important postlarval descriptions are included.

Trombiciila alfreddugesi has been the object of some concen-
trated studies. Most recently, Loomis (1956:1259-1280) presented
evidence on the factors influencing the abundance of larval T.
alfreddugesi on hosts. Some discussion was devoted to the post-
larval habitat. Evidently postlarvae normally inhabit grassy soil,
particularly in the more open areas. Although most trombiculids
have not been collected as postlarvae, nymphs and adults of T.
alfreddugesi are frequently found beneath limestone rocks and in
soil samples during the spring and early summer. Apparently they
overwinter as postlarvae.

This species is not strikingly distinct from the two other species
of Tromhicula (Eutromhicula) found in eastern Kansas. However,
by characters related to their smaller size, nymphs of T. alfred-

Nymphs of the Trombiculidae 247

dugesi are more readily separable from T. lipovskyana and T. splen-
dens than those two species are separable from each other.

Tromhicula alfreddugesi is a very common parasite upon many
species of both birds and mammals, and is found throughout most
of the United States. Among the nymphs available for study,
specimens from western Kansas and northern Texas were sig-
nificantly smaller in the measurement ASL than specimens from
eastern Kansas. No other differences were noted between speci-
mens from these areas.

Specimens examined. — Total 159 nymphs, as follows: Kansas.
Anderson County: 3 mi. N Garnett, Crotaphyttis collaris, July 8,

1948 (7 nymphs, reared LJL). Barber County: 4 mi. S Aetna,
Neofoma micropus, August 22, 1949 (39 nymphs, reared LJL),
July 25, 1952 (2 nymphs, reared DAG), and July 11, 1955 (15
nymphs, reared DAG, idet. by larval skins), and Sylvilagus fori-
danus, September 14, 1953 (7 nymphs, reared DAG); 5 mi. S Sun
Gity, Neotoma micropus, September 14, 1948 (2 nymphs, reared
LJL); 3 mi. S Aetna, Neotoma micropus, July 25, 1952 (2 nymphs,
reared DAG); 10^2 mi. W Hardtner, Sylvilagus floridanus, July 25,
1952 (2 nymphs, reared DAG). Cheyenne County: 3 mi. N St.
Francis, Perognathus hispidus, July 23, 1948 (1 nymph, reared
LJL), Pituophis catenifer, July 22, 1948 (1 nymph, reared
LJL). Douglas County: Univ. Kansas Natural History Reserva-
tion, Elaphe obsoleta, August 10, 1953 (12 nymphs, reared DAG,
idet. by larval skins ) , Crotaphytus collaris, June 24, 1950 ( 6 nymphs,
reared LJL), — "soil sample," February 27, 1951 (1 nymph, col.
RBL); 3 mi. S, 1 mi. E Lawrence, Sylvilagus floridanus, July 14,

1949 (1 nymph, reared LJL); 5 mi. S Lawrence, Sylvilagus flori-
danus, July 14, 1949 (7 nymphs, reared LJL); 4 mi. S Lawrence,
Zenaidura macroura, July 25, 1948 (1 nymph, reared LJL) and
Colinus virginiamis, July 25, 1948 (4 nymphs, reared LJL); 2 mi.
W Lawrence, Sylvilagus floridanus, June 19, 1949 (9 nymphs,
reared LJL). Miami County: Miami Go. State Park, Agkistrodon
contortrix, October 12, 1948 (1 nymph, reared LJL). Rawlins
County: 8 mi. S Beardsley, Perognathus hispidus, July 29, 1948
(1 nymph, reared LJL), and Cnemidophorus sexlineatus, July 29,
1948 (1 nymph, reared LJL). Seward County: Liberal, Dipodomys
ordii, September 8, 1948 (1 nymph, reared LJL); 4 mi. NE Liberal,
Sigmodon hispidus, September 12, 1948 (24 nymphs, reared LJL);
12 mi. NE Liberal, Sceloporus undulatus, September 9, 1948 (1
nymph, reared LJL). Shawnee County: 3 mi. W Topeka, Sylvila-
gus floridanus, August 26, 1948 (3 nymphs, reared LJL). Ne-

248 The University Science Bltlletin

BRASKA. Webster County: 3 mi. E Guide Rock, Heterodon nasicus,
August 5, 1951 (3 nymphs, reared DAC). Oklahoma. Woods
County: 6 mi. S, 2 mi. W Aetna, Kans., Heterodon platyrhinos,
October 7, 1951 ( 1 nymph, reared DAC ) . Texas. Taylor County:
Phrynosorna cornutum, June 10, 1952 (4 nymphs, reared DAC).

Trombicula helkini Gould
(Figs, 23, 79, 229)

Trombicula (Eutrombicula) belkini Gould, 1950, Wasmann Joiir. Biol., vol.
8, p. 367 (larva).

Description of nymph. — Resembles T. alfreddugesi except as fol-
lows:

Body: About 0.7 mm. long, color red. Eyes red, but obscured
in mounted materials, detectable by disruption of cuticular striae;
about 20 [x in diameter. Setal stem thickened, with many fine
branches; tips of posterior setae rounded, slightly expanded, with
numerous short branches; posterior setae about 65 \}. long. Sternum
with about 6 setae. Measurements of tarsus I (means of 6 speci-
mens); TL: 124.5 ±1.727, TH: 64.7 ± 0.4216, TL/TH: 1.92
± 0.0308.

Gnathosoma: Cheliceral blade with about 14 teeth. Cheliceral
measurements (means of 9 specimens); BL: 114.9 ± 0.8071, BH:
45.1 ± 0.7349, CL: 59.9 ± 0.4233, BL/BH: 2.57 ± 0.0500, BL/CL:
1.92 ± 0.0147. Palpus with dorsal femoral setae profusely branched;
tarsala about 9 \}. long. Setal counts; femur (7 specimens) : 7.21 =b
0.2641, genu (8 specimens): 11.00 ± 0.2631, tibia: range 5-6,
tarsus: range 8-10.

Scutum: Sensilla with branches shorter and more numerous;
bipectinate on distal portions. Scutal measurements (means of 6
specimens); ASL: 92.8 ± 0.7492, SB: 34.3 ±: 0.6062, TS: 33.7
± 0.9189, SENS: about 100, ASL SB: 2.73 ± 0.0616.

Remarks. — This species is readily separable from other species
of Trombicula (Eutromhicida) in the nymphal stage by the form of
the posterior body setae. Also, the profuse branching of the setae
on tlie palpal femur is a useable character. The eyes are of interest
in T. helkini; although they were detected in living material by
their color, the actual lens was visible on but one mounted speci-
men. The striae of the cuticle were disrupted in the area ad-
jacent to the sensillary bases, where the lens should be.

Nymphs of the Trombiculidae 249

Specimens examined. — Total 9 nymphs, as follows: California.
Marin County: Sceloporus occidentalis, June 1, 1952 (9 nymphs,
reared DAC).

Trombicula lipovskijana Wolfenbarger

(Figs. 25, 76)

Trombicula (Eutrombictila) lipovskijana Wolfenbarger, 1953, Ann. Ent. Soc.
America, vol. 45, pp. 660-666 (larva, postlarva).

Description of nymph. — Resembles T. alfreddugesi except as fol-
lows :

Body: About 1.0 mm. long, color red. Eyes sometimes obscured,
about 20 [J. in diameter. Posterior body setae slightly shorter,
about 50 [J. long. Sternum with about 12 setae. Measurements
of tarsus I (means of 8 specimens); TL: 149.0 ± 3.742, TH:

81.3 ± 2.102, TL/TH: 1.85 ±: 0.0191.

Gnathosoma: Cheliceral blade with about 22 teeth. Cheliceral
measurements (means of 10 specimens); BL: 140.0 rt 2.936, BH:

61.4 ± 0.8326, CL: 84.4 ± 1.0872, BL/BH: 2.32 ± 0.0359, BL/CL:
1.66 ± 0.0276. Palpus large; tarsala about 13 \s. long. Setal
counts; femur: 6.5 ± 0.2835 (8 specimens), genu: 14.05 d= 0.3877
(10 specimens), tibia: range 4-6, tarsus: range 8-10.

Scutum: Measurements (means of 10 specimens); ASL: 116.9 rt
1.656, SB: 47.9 ± 0.8360, TS: 35.9 ± 2.0911, SENS: about 140,
ASL/SB: 2.44 ± 0.0400.

Remarks. — As larva, this species is very close to T. alfreddugesi,
but as nymph its affinities appear to be with T. splendens. Indeed
T. lipovskyana may be a subspecies of T. splendens. The distribu-
tion of T. lipovskyana is not well known; apparently it is distributed
throughout the southeastern United States (see Loomis, 1956:1280),
The habitat of the postlarvae is similar to that of T. alfreddugesi,
being grassy soil.

Specimens examined. — Total 24 nymphs, as follows: Kansas.
Douglas County: 3 mi. S, 1 mi. E Lawrence, Sylvilagus floridanus,
July 14, 1949 (11 nymphs, reared LJL); 2 mi. W Lawrence, Sylvila-
gus floridanus, June 19, 1949 (7 nymphs, reared LJL), Louisiana,
Jefferson Parish: Anolis carolinensis, April 28, 1954 (2 nymphs,
reared RBL-DAC, idet, by larval skins). Orleans Parish: 15 mi.
NE New Orleans, Anolis carolinensis, April 27, 1954 (4 nymphs,
reared RBL, 2 idet, by larval skins).

250 The University Science Bulletin

Trombictila splendens Ewing
(Figs. 13-15, 24, 75, 176, 199, 228)

Trombicula splendens Ewing, 1913, Bull. Amer. Mus. Nat. Hist., vol. 32, pp.

113-114 (adult).
Trombicula (Eutrombicula) splendens, Jenkins, 1949, Ann. Ent. Soc. America,

vol. 42, p. 304 (adult); Wolfenbarger, 1953, Ann. Ent. Soc. America, vol.

45, p. 650 (adult).
Trombicula (Trombicula) splendens, Womersley, 1952, Rec. South Australian

Mus., vol. 10, p. 334 (adult).

Description of nymph. — Resembles T. alfreddugesi except as fol-
lows :

Body: About 1.0 mm. long, color red (rarely white). Eyes about
20 [X in diameter. Tips of posterior setae usually with 3 short
branches, rarely with 2; posterior setae about 60 [jl long. Measure-
ments of tarsus I (means of 9 specimens); TL: 142.8 ± 1.793,
TH: 77.1 ± 1.148, TLTH: 1.83 ± 0.167.

Gnathosoma: Cheliceral blade with about 20 teeth. Cheliceral
measurements (means of 10 specimens); BL: 140.8 ± 1.332, BH
55.9 ± 0.5859, CL: 77.8 ± 0.7859, BL/BH: 2.51 ± 0.0348, BL/CL
1.81 ± 0.0179. Palpus large, tarsala about 14 [i. long. Setal counts
femur (9 specimens) : 5.06 ± 0.2324, genu (3 specimens) : 12.88 ±
0.5774, tibia: range 5-7, tarsus: range 8-10.

Scutum: Measurements (means of 10 specimens); ASL: 114.6 ±
2.202, SB: 45.3 ±2.791, TS: 44.6 d= 1.248, SENS: about 140,
ASL/SB: 2.53 ±0.0298.

Remarks. — As in the case of T. alfreddugesi, the foregoing syn-
onomy contains only the more important references to postlarvae
of the species. Womersley 's description of the sensilla as nude is
at variance with my findings.

This species is known from Kansas from adults only. Adults
taken on several occasions in decaying logs in Miami County were
identified by larvae obtained in culture as T. splendens. Most of
the nymphs here described are second generation, being reared
in such a culture of materials from Titus County, Texas.

Tromhicida splendens was originally described from adults. The
proper larval form was associated by Jenkins (1949:201).

Specimens examined. — Total 20 nymphs, as follows: Louisiana.
St. Charles Parish: 17 mi. SW New Orleans, Agkistrodon piscivor-
ous, May 1, 1954 (1 nymph, reared RBL, idet. by larval skin).

Nymphs of the Trombiculidae 251

Texas. Titus County: 3 mi. NW Talco, March 26, 1951 (Adults
taken from decaying logs; second generation nymphs obtained;
total 19).

Subgenus Neotrombicula Hirst

Neotrombicula Hirst, 1925, Nature, vol. 116, p. 609 (type Acarus autumnalis

Shaw ) (larva).
TTombicula (Neotrombicula), Brennan and Wharton, 1950, American Midi.

Nat., vol. 44, p. 156 (adult); Philip and Fuller, 1950, Parasit., vol. 40, pp.

54-55 (posthrvae); Womersley, 1952, Rec, South Australian Mus., vol. 10,

p. 349 (adult).

Diagnosis (nymph). — None.

Remarks. — I find no character or group of characters which permit
recognition of this subgenus. Therefore it is left without diagnosis
in the nymphal stage, although larvae can be recognized.

Certain features of possible subgeneric importance should be
mentioned. The four species studied by me have stout cheliceral
bases and wide blades, with rather large teeth. The least extreme
is T. autumnalis, which has chelicerae more similar to those of
Trombicula (Eutrombicula). Sensilla tend to be thickened and
provided with small branches (except T. sylvilagi). In these fea-
tures the species resemble some members of Euschongastia more
closely than they resemble other species of Trombicula. It is in-
teresting to note that in both genera the species so modified are
fall and winter chiggers. A better source of group characters for
these species appears to be the scutum. The species all have
similar scuta but I have not been able to find a character which
separates them from nymphs of other Trombicula species. A very
possible character involves the bulla of the sensillary area, which
is somewhat enlarged and either lacks a carina or has a short, thick
carina almost indistinguishable from the bulla itself. This single
feature would not exclude some other species of Trombicula from
the subgenus Neatrombicula.

Nymphs of at least three species of the subgenus Neotrombicula
have been previously described. Trombicula pomeranzevi Schluger
has been described by Sasa. Andre has described the nymph of
T. autumnalis. Several descriptions are available of postlarvae of
T. japonica Tanaka but tliese are inconsistent; the nymph described
by Womersley (1952:318, after Tanaka) is certainly not the same
as that described by Sasa (1953:423), as noted by Sasa.

252 The University' Science Bulletin

Tromhiculo autumnahs (Shaw)
(Figs. 52, 82, 157, 236)

Acants autumnolis Shaw, 1790, Naturae vivarii (Nat. Misc.), vol. 2 (no pag-
ination ) (larva).

Trombicula auttimnalis, Kneissl, 1916, Zool. Anz., vol. 46, p. 253 (nymph);
Hirst, 1915, Jour. Econ. Biol., vol. 10, p. 73 (nymph); Philip and Fuller,
1950, Parasit., vol. 40, p. 55 (nymph); Richards, 1950, Parasit., vol. 40, pp.
105, 115 (postlarvae); Jones, 1951, Parasit., vol. 41, pp. 241-245 (postJarvae);
Womersley, 1952, Rec. South Australian Mus., vol. 10, p. 364 (postlarvae).

Thromhicula autumnalis, Andre, 1929, Bull. Must. d'Hist. Nat. Paris, 2e Ser.,
vol. 1, p. 394 (adult); Andre, 1930, Mem. Zool. Soc. France, vol. 29, pp.
39-138 (postlarvae); Andre, 1937, Bull. Mus. d'Hist. Nat. Paris, vol. 9, p.
313 (adult).

Description of nymph. — Body: Elongated, about 0.7 mm. long,
color not known. Setae normal, with long branches; tips com-
posed of 1 or 2 somewhat long branches; posterior setae about 70 [x
long. Sternum roughly pentagonal, closed behind, with about 10
setae. Measurements of tarsus I (means of 9 specimens); TL:
154 ±: 5.077, TH: 71.4 ± 1.271, TL/TH: 2.13 ± 0.0441.

Gnathosoma: Chelicera nearly of usual shape, perhaps slightly
robust; base punctate; blade with about 15 teeth. Cheliceral meas-
urements (means of 10 specimens); BL: 122.4 ± 2.509, BH:
49.2 ± 0.8138, CL: 70.5 ± 1.014, BL/BH: 2.51 ± 0.0568, BL/CL:
1.73 ± 0.0213. Palpus somewhat stout, very similar to that of T.
lipovshji; tarsala about 11 [x long; tarsus with 3 inner, 2 outer
apical nude setae. Setal counts (means of 10 specimens); femur:-
5.17 ± 0.1179, genu: 8.3 =iz 0.2135, tibia: range 4-5, tarsus: range
9-10.

Scutum: Tectum broad, with about 12 teeth on margin, very
irregular. Sensillary area broadly triangular in shape; basal plate
of two widely separated bands; apodeme conspicuous; strong ridges
anterior and posterior to sensillary bases; ridges medial to sensillary
bases very weak, posterior ridge very weak; bulla triangular, short
and broad, punctate; carina not seen; sensillary bases rather small;
sensilla somewhat thickened, with short branches on basal thirds
and longer ones on distal portions. Scutal measurements (means
of 9 specimens); ASL: 97.0 ± 4.384, SB: 44.3 ± 1.476, TS: 34.9
± 1.493, SENS: about 115, ASL/SB: 2.19 zt 0.0423.

Remarks. — I am indebted to Mr. D. M. Minter for these speci-
mens of T. autumnalis.

Trombicula autumnalis is probably the species best known as
postlarva, thanks to the careful works of Andre (particularly the
1930 paper), Richards (1950), and Jones (1951).

Nymphs of the Trombiculidae 253

Specimens examined. — Total 10 nymphs, as follows: England.
Kent: Chevenins Park, August-September, 1954, "wild rabbits,"
(10 nymphs, reared by D. M. Minter; associated with larvae of
normal type "B" of Richards).

Trombicula lipovskyi Brennan and Wharton
(Figs. 47, 83, 116, 117, 158, 240)

Trombicula (Ncotrombicula) lipovskyi Brennan and Wharton, 1950, American
Midi. Nat., vol. 44, p. 177 (larva); Kardos, 1954, Univ. Kansas Sci. Bull.,
vol. 36, pp. 91-98 (larva).

Description of nymph. — Body: Rounded, about 0.6 mm. long,
color yellow to orange. Setae with stems slightly thickened,
branches fine; tips of posterior setae rounded, with about 3 rather
long branches; posterior setae about 45 [jl long. Sternum pentag-
onal, closed behind, with about 6-8 setae. Measurements of tarsus
I (means of 10 specimens); TL: 128.9 ± 3.971, TH: 64.6 ± 1.628,
TL/TH: 2.00 ±0.0259.

Gnathosoma: Chelicera with both base and blade robust; base
punctate, blade with about 11 teeth of intermediate size. Cheliceral
measurements (means of 39 specimens); BL: 117.1 ± 1.135, TH:
48.9 zh 0.4778, CL: 64.3 ± 0.4776, BL/BH: 2.41 ± 0.0275, BL/CL:
1.83 ± 0.0172. Palpus of usual shape; claw shghtly elongated; tar-
sala about 11 pi long; tarsus with 3 inner, 2 outer apical nude setae.
Setal counts (means of 10 specimens); femur: 4.00 ± 0.4887, genu:
8.2 ± 0.3092, tibia: range 3-5, tarsus: range 8-10.

Scutum: Tectum triangular in shape, margin with about 10
teeth. Sensillary area markedly triangular in shape; apodeme con-
spicuous; ridges rather strong but usually distorted; bulla trian-
gular, prominent, apparently highly arched; carina not seen; sensil-
lary bases of usual size; sensilla somewhat thickened, flexible, with
short branches distributed along entire length, with few longer
branches near tips; stems often but not always furcate at tips.
Scutal measurements (means of 10 specimens); ASL: 83.8 ± 1.373,
SB: 38.1 ± 0.994, TS: 36.5 ± 1.046, SENS: about 110, ASL/SB:
2.21 ± 0.0586.

Remarks. — Trombicula lipovskyi was reared to the adult stage.
Nymphs ate Collembola eggs.

This species is a common chigger in the eastern part of the Mid-
west Region. Its closest relative in both larval and nymphal stages
appears to be T. whartoni.

Specimens examined. — Total 153 nymphs, as follows: Kansas.
Barber County: 3/2 mi. S, 1 mi. W Aetna, Neotoma micropus, April

254 The University Science Bulletin

11, 1949 (12 nymphs, reared LJL). Douglas County: Lawrence,
Microtus ochrogaster, March 5, 1949 (1 nymph, reared LJL),
— Sigmodon hispidus, October 15, 1948 (1 nymph, reared LJL),
and Neotoma floridana, March 30, 1949 (4 nymphs, reared LJL);
3 mi. W Lawrence, Neotoma floridana, March 5, 1949 (8 nymphs,
reared LJL) — Sylvilagus floridanus, November 12, 1949 (46 nymphs,
reared LJL); 9 mi. W Lawrence, Sylvilagus floridanus, January

2, 1949 (6 nymphs, reared LJL); 4 mi. N, 1 mi. E Lawrence,
Neotoma floridana, November 26, 1948 (9 nymphs, reared LJL);
2 mi. S, 2 mi. W Pleasant Grove, Sylvilagus floridanus, March

3, 1949 (1 nymph, reared LJL), December 4, 1948 ( 1 nymph, reared
LJL), and December 29, 1948 (1 nymph, reared LJL); 3 mi. S
Eudora, Sylvilagus floridanus, February 17, 1949 ( 1 nymph, reared
LJL); 3 mi. W, 2 mi. S CHnton, Sylvilagus floridanus, March 10,
1949 (1 nymph, reared LJL). Jefl^erson County: 6 mi. E Perry,
Neotoma floridana, December 28, 1948 (3 nymphs, reared LJL);
5/2 mi. N, /2 mi. E Lawrence, Sylvilagus floridanus, November 21,
1951 (17 nymphs, reared DAC), and November 24, 1951 (4 nymphs,
reared DAC). Johnson County: 2 mi. N, 1 mi. W Lenexa, Syl-
vilagus floridanus, November 18, 1953 (13 nymphs, reared RBL).
Wyandotte County: Kansas City, Sylvilagus floridanus, October 30,
1953 (1 nymph, reared RBL), and December 21, 1953 (3 nymphs,
reared RBL). Oklahoma. Woods Cowity: 6 mi. S, 2 mi. W Aetna,
Kans., Heterodon platyrhinos, October 7, 1951 (1 nymph, reared
DAC).

Trombicula sylvilagi Brennan and Wharton

(Figs. 53, 160, 237)

Trombicula (Neotrombicula) sylvilagi Brennan and Wharton, 1950, Amer.
Midi. Nat., vol. 44, pp. 186-187 (larva); Kardos, 1954, Univ. Kansas Sci.
Bull., vol. 36, pp. 103-111 (larva).

Description of nymph — Resembles T. lipovskyi except as follows:
Body: Elongated, small (0.5 mm. long), color orange. Setae
short, stems thick; tips rounded, with 2-3 short branches; posterior
setae about 25 [x long. Measurements of tarsus I ( means of 5 speci-
mens); TL: 101.4 ±0.600, TH: 55.0 ± 0.632, TL/TH: 1.84 ±
0.0283.

Gnathosoma: Cheliceral base with conspicuously large puncta;
blade with about 10 particularly large teeth. Cheliceral measure-
ments (means of 4 specimens); BL: 85.3 ± 1.315, BH: 37.5 ± 0.866,
CL: 52.5 db 1.255, BL/BH: 2.25 ± 0.0646, BL/CL: 1.63 ± 0.0629.
Palpus with tarsala about 9 [i long. Setal counts; femur (1 speci-
men): 3.5, genu (3 specimens): 6.50 ±: 0.2887, tibia: range 4-5,
tarsus: 9.

Nymphs of the Trombiculidae 255

Scutum: Sensilla with short branches on basal thirds, longer ones
on distal portions (resembling T. autumnalis), stems slightly ex-
panded distally. Scutal measurements (means of 5 specimens);
ASL: 65.4 ± 1.0296, SB: 34.2 ± 0.800, TS: 14.3 ± 0.4787, SENS:
about 75, ASL/SB: 1.92 ± 0.0490.

Remarks. — As a larva, this species was listed by Brennan and
Wharton as "ungrouped," since it did not appear closely related to
other species in Neotrombicula. As a nymph, T. sylvilagi is not
markedly diflFerent from T. lipovskiji, except in being smaller. Until
more postlarvae of Neotrombicula are known, it is not possible to
evaluate "groups" of species in 'Neotrombicula except on larval
characters.

The specimens studied were reared by Ervin H. Kardos in an
unusual manner. Unengorged larvae were obtained with chigger
samplers and were then allowed to attach to a laboratory mammal
( Mus ) . Nymphs were then obtained from these larvae.

Specimens examined. — Total 5 nymphs, as follows: Kansas.
Douglas County: Univ. Kansas Natural History Reservation, chigger
sampler, October 8, 1952 (5 nymphs, reared EHK).

Trombicula whartoni Ewing

(Figs. 51, 80, 159)

Trombicula whartoni Ewing, 1929, Ent. News, vol. 40, p. 296 (larva).
Trombicula ( Neotrombicula ) whartoni, Brennan and Wharton, 1950, Amer.

Midi. Nat., vol. 44, pp. 175-176 (larva); Kardos, 1954, Univ. Kansas Sci.

Bull., vol. 36, pp. 98-102 (larva).

Description of nymph. — Resembles T. lipovskyi except as follows:

Body: About 0.7 mm. long. Measurements of tarsus I (means of
9 specimens); TL: 148.2 ± 2.101, TH: 77.9 ± 1.012, TL/TH:
1.91 ± 0.0247.

Gnathosoma: Chelicerae nearly normal in shape, perhaps slightly
robust; base with conspicuous puncta; blade with about 15 teeth of
intermediate size. Cheliceral measurements (means of 9 speci-
mens); BL: 120.6 ±: 1.925, BH: 47.1 ± 1.181, CL: 65.9 ± 0.8233,
BL/BH: 2.58 ± 0.0843, BL/CL: 1.83 =b 0.0211. Palpus with tarsala
about 11 [JL long; tar.sus with 3 inner, 2-3 outer apical nude setae.
Setal counts; femur (6 specimens); 6.00 ± 0.3874, genu (9 speci-
mens): 10.17 ± 0.4714, tibia: range 4-6, tarsus: range 10-15.

Scutum: Sensillary area with bulla with conspicuous puncta;
sensilla nude except for small basal branches, rarely furcate at tips.
Scutal measurements (means of 9 specimens); ASL: 101.9 ±
1.775, SB: 42.8 ± 1.038, TS: 40.3 ± 1.291, SENS: about 145,
ASL/SB: 2.41 ± 0.0484.

256 The University Science Bulletin

Remarks. — The nymph is similar to that of T. lipovshji but is
easily differentiated by its nude sensilla. Loomis (1956:1295) sug-
gests that T. whartoni seems restricted to deciduous woods and
the woodland edge, while T. lipovshji occurs in the drier open grass-
lands but also in the woodland edge.

Trombictda whartoni was less frequently collected than T. lip-
ovskyi, but the difference is not so great as to explain why no T.
whartoni nymphs appeared in the mixed cultures maintained by
L. J. Lipovsky. On the two occasions when the species was suc-
cessfully cultured, the cultures were pure.

Specimens examined. — Total 9 nymphs, as follows: Kansas.
Douglas County: 2 mi, S Warden, Sciurus niger, November 28,
1949 (2 nymphs, reared LJL). Johnson County: Roeland Park,
Sylvilagus floridanus, November 4, 1953 (7 nymphs, 1 idet. by lar-
val skin, reared RBL-DAC).

Subgenus Leptotromhidiiim Nagayo, Miyagawa, Mitamura

and Imamura

Leptotrombidium Nagayo et al., 1916, Dobutsugaku Zasshi, vol. 28, p. 392
(type Trombidiutn akamushi Bninipt) (larva).

Trombicula (Leptotromhidiiim), Ewing, 1949, J. Washington Acad. Sci., vol.
39, p. 236 (adidt); Womersley, 1952, Rec. South AustraHan Mus., vol. 10,
p. 326 (aduJt); Sa.sa, 1953, Japanese Jour. Exp. Med., vol. 23, p. 411 (post-
larvae); Audy, 1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 140
(postlarvac).

Diagnosis (nymph). — (2) Precoxal plates of coxae I present but
fused in midline to form longitudinally divided sternum.

Remarks. — This diagnostic feature for Leptotrombidium was first
used by Womersley; Sasa and Audy have followed his diagnosis
and the nymphs described by Sasa agree with it. The nymph of
T. myotis, described in the present paper, is the seventeenth species
of T. (Leptotrombidium) to be knovra in the postlarva stage.

Prior to Womersley's (1952:326) discovery of the divided ster-
num as a subgeneric feature, the subgenus Leptotrombidium was
diagnosed on combinations of larval characters. It seems likely
that in the future, workers will place more emphasis on the nymphal
stage for recognition of this subgenus. Leptotrombidium was ex-
panded by Womersley (1952:326) and further expansion was sug-
gested by Audy (1954:140). The limits of the subgenus are cer-
tainly vague at present; it seems doubtful that the single diagnostic
feature used for the postlarval stages will stand the test of time.

Nymphs of the Trombiculidae 257

Trombicula mijotis Ewing
(Figs. 55, 81, 106, 107, 166. 231)

Trombicula myotis Ewing, 1929, Ent. News, vol. 40, pp. 294-295 (larva).
Trombicula (Leptotrombidium) myotis, Wharton and Fuller, 1952, Mem. Ent.

Soc. Washington, no. 4, p. 54 (larva); Loomis, 1956, Univ. Kansas Sci. Bull.,

vol. 37, p. 1287-1288 (larva).

Description of mjmph. — Body: Rounded posteriorly, about 0.6
mm. long, color not known. Setae long, branches fine; tips of
posterior setae spatulate, with several short branches; posterior
setae about 60 [i long. Sternal division rarely incomplete; each
side with about 4 setae. Measurements of tarsus I (means of 4
specimens); TL: 101.3 + 0.4787, TH: 59.8 ± 0.2500, TL/TH:
1.7 ± 0.000.

Gnathosoma: Cheliceral base somewhat robust, punctate, area
of articulation large; blade shearlike, apodeme of blade not con-
stricted, with about 19 small teeth. Cheliceral measurements
(means of 4 specimens); BL: 93.0 ± 1.528, BH: 37.8 ± 2.462,
CL: 49.8 ±: 1.250, BL/BH: 2.63 ± 0.0885, BL/CL: 1.83 ± 0.0408.
Palpus of usual shape; tarsala about 11 \}. long, claw rather short;
tarsus with 3 inner, 1 outer apical nude setae. Setal counts ( means
of 4 specimens); femur: 4.88 ± 0.0126, genu: 7.00 ± 0.4787, tibia;
5, tarsus: 9.

Scutum: Tectum large, square, with about 10 teeth. Crista nar-
row. Sensillary area broad, oval; basal plate of two rather large
halves; apodeme short; strong ridges anterior and posterior to
sensillary bases, others very weak; bulla much reduced, triangular;
carina broad but short; sensillary bases prominent; sensiUa some-
what thickened but flexible, with short branches on basal half,
longer ones on distal half, longer branches fine. Scutal measure-
ments (means of 3 specimens); ASL: 80.3 d= 2.028, SB: 31.3 ±
0.8539, TS: 29.3 ± 0.3334, SENS: about 80, ASL/SB: 2.50 ±
0.0577.

Remarks. — Loomis (1956:1289) suggests that postlarvae of this
species are probably inhabitants of decaying wood, and possibly
nests of mammals nesting in such places.

The nymph of T. myotis cannot be keyed down to species of T.
(Leptotrombidium) in the key given by Womersley (1952:342).
Sasa (1953:411) experienced diflBculty in separating his nymphs
of Japanese species. I am unable to separate T. myotis from the
nymphs described by Sasa. It appears that the subgenus Lep-

9—8920

258 The University Science Bulletin

totrombidium is a group of species far more readily separable in the
larval stage than in postlarval stages.

Specimens examined. — Total 4 nymphs, as follows: Iowa. Fre-
mont County: Waubonsie State Park, 5 mi. S, 2 mi. W. Sidney,
Feromij&cus leucopus, October 10, 1953 (4 nymphs, reared RBL).

Subgenus Trombicula Berlese

Trombicula Berlese, 1905, Redia, vol. 2, p. 155 (type Trombicula minor Ber-
lese ) ( adult ) .

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates ab-
sent. (10) Tectum square to triangular, with few to many teeth on
margin. (22) Sensillary area with posterior scutal ridge without
elevations or with only weak elevations. ( 23 ) About eight to twelve
parascutal setae.

Remarks. — More realistically, this subgenus contains those species
which do not fall into the other subgenera. The impossibility of
recognizing T. minor has been previously discussed; without a usa-
ble type species the subgenus can only serve as a catch-all.

About twenty postlarvae which would apparently fall into this
subgenus have been described; about ten of these are known as
adults only. Inclusion of these forms in the key to species was
not feasible.

Trombicula crossleyi Loomis

(Figs. 132, 133, 163)

Trombicula crossleyi Loomis, 1954, Univ. Kansas Sci. Bull., vol. 36, pp. 920-
922 (larva).

Description of nymph. — Body: Small (about 0.5 mm. long),
elongated, color orange to red. Setae somewhat short, stem rather
thickened, branches fine and numerous; tips of posterior setae with
about 3-5 short branches; posterior setae about 20 [x long. Sternal
area elongated posteriorly, rectangular in shape, usually open
behind, with about 4 setae. Measurements of tarsus I ( means of 10
specimens); TL: 77.0 ± 0.7889, TH: 36.8 ± 0.3267, TL/TH: 2.10
± 0.0298.

Gnathosoma: Cheliceral base of usual shape, puncta almost
absent; blade shearlike, with about 14 rather small teeth. Cheliceral
measurements (means of 6 specimens); BL: 81.6 ± 1.70, BH:
28.4 ± 0.68, CL: 36.8 ± 0.76, BL/BH: 2.94 ± 0.093, BL/CL: 2.26
± 0.034. Palpus slightly elongated; dorsal genual setae often nude,
claw and accessory claws rather small, tarsala about 6 [x long; tarsus
with 2 inner, 1 outer apical nude setae. Setal counts (means of 6
specimens); femur: 3.00 (no variation), genu: 2.83 ± 0.1258; tibia:
range 4-5, tarsus: range 7-9.

Nymphs of the Trombiculidae 259

Scutiim: Tectal margin with about 10 conspicuous teeth. Sen-
sillary area nearly transverse-oval in appearance; apodeme reduced;
ridges weak, usually distorted; bulla triangular; carina small and
thin; sensillary bases prominent; a pair of noticeably elongated para-
scutal setae directly in front of sensillary area; sensilla straight,
slightly expanded distally, branches short on basal half, longer on
distal half. Scutal measurements (means of 10 specimens); ASL:
59.2 ± 1.083, SB: 27.3 ± 1.041, TS: 20.8 ± 0.249, SENS; about 65,
ASL/SB: 2.18 ± 0.1065.

Remarks. — This species is one of a group which, as nymphs, pos-
sesses several key characters apparently associated with somewhat
smaller size than most species of Trombicida. This group of species
contains T. crossleiji, T. trisetica, T. merrihewi, and T. ornata; some
features distinguishing this group are the presence of but three
apical nude setae on the palpal tarsus, usually but three setae on
the palpal femur, prominent sensillary bases on the scutum (seem-
ingly a reduction of scutal size rather than enlargement of the sen-
sillary bases), and a tendency towards elongation of one pair of
parascutal setae. The larvae of these four species are more diverse;
larvae of T. crossleiji and T. trisetica are similar and are undis-
tinguished Trombictila species but larvae of T. merrihewi and T.
ornata, while resembling each other, have distinctive scuta and
chelicerae.

It is interesting to note that these four species, although similar
in morphology in postlarval stages, seemingly exist in diflFerent
habitats. Loomis (1956:1322), on the basis of larval host prefer-
ences, suggests the habitat of postlarvae of T. crossleiji and T.
trisetica to be standing dead trees, and that of T. ornata to be
crevices of rock outcroppings and possibly nests of mammals, while
T. merrihewi is known only from bats and postlarvae are assumed
to be cave dwellers. Thus it seems likely that the features common
to these species are associated with small size rather than being
direct adaptations to specific habitats.

Of the species placed in the subgenus Trombicula, these four
appear to be the most likely relatives of Trombicula minor. Tin's
relationship is suggested not only by their small size but by com-
parison of their scuta with the illustration of that of T. minor, as
given by Willmann (1940:133). As previously noted, Willmann's
drawing is diagrammatic and may not be adequate for such com-
parison.

Trombicula crossleyi is very similar to T. trisetica in both larval
and postlarval stages, and eventually the two may prove to be

260 The University Science Bulletin

conspecific. Nymphs are difficult to separate; the only differences
found involve measurements.

Specimens examined. — Total 12 nymphs, as follows: Kansas.
Barber County: lOM mi. W Hardtner, Melanerpes erythrocephalus,
July 26, 1952 (5 nymphs, reared DAC); 4 mi. S Aetna, Feromyscus
leucopus, October 7, 1951 (6 nymphs, reared LJL), July 25, 1952
( 1 nymph, reared DAC ) .

Trombictila fitchi Loomis
(Fig. 230)

Trombicula fitchi Loomis, 1954, Univ. Kansas Sci. Bull., vol. 36, pp. 926-928

(larva).
Trombicula (Neotrombicula) fitchi, Loomis, 1956, Univ. Kansas Sci. Bull., vol.

37, pp. 1297-1298 (larva).

Description of nymph. — Body: Rounded, about 0.5 mm. long,
color not known. Setae with stems normal, branching distinctive;
posterior setae with long fine branches on basal halves but short
thick-based branches on distal halves; tips with 2 or 3 short branches;
posterior setae about 60 \i long. Sternal area roughly pentagonal
in shape, closed behind, with about 8 setae. Measurements of
tarsus I (means of 10 specimens); TL: 109 ± 3.157, TH: 53.5 ±
1.046, TL/TH: 2.03 ± 0.0423.

Gnathosoma: Chelicerae of usual shape, base punctate; blade
with about 20 rather small teeth. Cheliceral measurements (means
of 113 specimens); BL: 111.6 ± 1.819, BH: 38.4 ± 1.071, CL:
55.5 ± 1.096, BL/BH: 2.92 ± 0.0715, BL/CL: 2.01 ± 0.0265. Palpi
of usual shape, dorsal genual setae occasionally nude; tarsus with
tarsala about 8 }jl long; tarsus with 3 inner, 2 outer apical nude
setae. Setal counts (means of 8 specimens); femur: 3:94 zb 0.2904,
genu: 8.13 ± 0.3239, tibia: range 2-5, tarsus: range 8-9.

Scutum: Tectum with about 8 teeth on margin. Sensillary area
triangular; basal plate of two bands; apodeme well developed;
ridges usually distorted but rather strong; bulla triangular, well
developed; carina thick, hard to see; sensilla ratlier thick but flex-
ible, with short branches basally but somewhat larger branches
distally; branches generally few, none large. Scutal measurements
(means of 10 specimens); ASL: 92.9 ± 1.853, SB: 35.3 ± 1.033,
TS: 33.7 ± 0.5588, SENS: about 125, ASL /SB: 2.63 ± 0.0472.

Remarks. — As nymph, Trombicula fitchi is easily recognized by
the form of the posterior body setae. Nymphs of T. kardosi have
similar posterior setae but the two are still sufficiently different to
permit easy separation.

Nymphs of the Trombiculidae 261

Loomis (1956:1290) placed these two species in the subgenus
Neotrombicula on the basis of larval characteristics. To do so re-
quired a broadening of the concept of the subgenus which elim-
inated the principal key character for the subgenus (the presence
of long nude whiplike setae on leg III). In the nymphal as well
as in the larval stage, T. fitchi and T. kardosi appear to me to be
related to species of T. (Neotrombicula), as Loomis suggests. How-
ever, it is difficult to determine how close this relationship may be.
Until more nymphs of species of T. (Neotrombicula) are known,
I prefer to retain T. fitchi and T. kardosi in the subgenus Trom-
bicula.

Specimens examined. — Total 17 nymphs, as follows: Kansas.
Barber County: 3/2 mi. S Aetna, Myotis velifer, April 10, 1949 (5
nymphs, reared LJL). Douglas County: Sciurus niger, January
19, 1950 (2 nymphs, reared LJL); 2 mi. S Warden, Sciurus niger,
November 26-28, 1949 (10 nymphs, reared LJL).

Trombicula gurneyi Ewing
(Figs. 57, 69, 112, 113, 164, 238)

Trombicula gurneyi Ewing, 1937, Proc, Biol. Soc. Washington, vol. 50, p.
169 (larva); Loomis, 1955, Univ. Kansas Sci. Bull., vol. 37, pp. 252-257
(larva).

Description of nymph. — Body: Rounded, about 0.6 mm. long,
color red to orange. Setae somewhat short, stems rather thickened,
branches fine and numerous; tips of posterior setae with about 3
short branches; posterior setae about 30 [x long. Sternum closed
behind, roughly pentagonal, with about 6 setae. Measurements
of tarsus I (means of 11 specimens); TL: 103.0 ± 3.555, TH:
53.4 ± 1.330, TL TH: 1.93 ± 0.0557.

Gnathosoma: Cheliceral base of usual shape, puncta few; blade
shearlike, with about 16 teeth. Chehceral measurements (means
of 8 specimens); BL: 94.1 ±: 1.865, BH: 36.0 ± 1.210, CL: 49.8
± 1.146, BL/BH: 2.63 ± 0.0525, BL/CL: 1.88 ± 0.0249. Palpus of
usual shape; tarsala about 11 [j. long; tarsus with 3 inner, 2 outer
apical nude setae. Setal counts (means of 9 specimens); femur:
4.94 ± 0.2693, genu: 8.50 ± 0.2635, tibia: range 4-5, tarsus: range
8-10.

Scutum: Tectum with about 8 rather small teeth. Sensillary area
triangular; basal plate of two widely separated bands; apodeme
prominent; bulla triangular; carina thin, long; ridges rather strong,
ridges medial to sensillary bases fused with posterior ridge and
somewhat prominent at this point (resembling members of the sub-

262 The University Science Bulletin

genus Eutrombicula); sensilla flagelliforin, flexible, stems thin and
finely tapered, with basal branches short and distal branches few,
fine, and long. Scutal measurements (means of 11 specimens);
ASL: 80.1 ± 2.605, SB: 37.4 ± 1.171, TS: 30.4 ± 1.974, SENS:
about 100, ASL/SB: 2.11 ± 0.0489.

Remarks. — Trombicula ^itrneyi is closely related to T. kansas-
ends, as seen in both larval and nymphal stages. The species may
be separated by the larger size and elongated cheliceral base of
T. kansasensis.

These two species show puzzling similarities to members of the
subgenus Eittrojubicula in the nymphal stage. Both species have
the fine flagelliform sensilla with fine branches and the "bilobed"
appearance of the posterior scutal ridge, although this latter feature
is not so pronounced as in true Eiitromhiciila.

Trombicula gurneiji is rather easily cultured and has been reared
through a complete generation in the laboratory. Food of the
postlarval stages was collembolan eggs.

Specimens examined. — Total 16 nymphs, as follows: Arkansas.
Little River County: 5 mi. S Ashdown, Eumeces laticeps, May 3,
1954 (4 nymphs, reared RBL-DAC). Kansas. Barber County:
Aetna, Dipodomys ordii, July 11, 1955 (2 nymphs, reared DAC);
2 mi. S Aetna, Dipodomys ordii, September 14, 1953 (1 nymph,
reared DAC, idet. by larval skin ) : 4 mi. S Aetna, Peromyscus manic-
ulatus, October 7, 1951 (1 nymph, reared LJL). Douglas County:
Univ. Kansas Natural History Reservation, Eumeces fasciatus, June
27, 1952 (2 nymphs, reared DAC), and Peromyscus maniculatus,
October 20, 1951 (3 nymphs, reared DAC). Rawlins County:
Citellus tridecemlineatus, July 28, 1948 (2 nyinphs, reared LJL).
Nebraska. Webster County: 3 mi. E Guide Rock, Heterodon
nasicus, August 5, 1951 (1 nymph, reared LJL).

Trombicula kansasensis Loomis

(Figs. 56, 77, 114, 115)

Trombicula kansasensis Loomis, 1955, Univ. Kansas Sci. Bull., vol. 37, pp.
260-262 (larva).

Description of nymph. — Resembles T. gurnet/i except as follows:

Body: About 0.8 mm. long, color orange. Setae similar to those

of T. gurneyi but longer; posterior setae about 65 [x long. Sternum

with 4 setae ( 1 specimen ) . Measurements of tarsus I ( 1 specimen ) ;

TL: 128, TH: 58, TL/TH: 2.2.

Nymphs of the Trombiculidae 263

Gnathosoma: Cheliceral base elongated, punctate, area of articu-
lation elongated; apodeme of blade strongly constricted; blade with
about 18 teeth. Cheliceral measurements (1 specimen); BL: 120,
BH: 40, CL: 62, BL/BH: 3.0, BL/CL: 1.9. Palpus large, rather
elongated, tarsala about 12 [l long. Setal counts (1 specimen);
femur: 6, genu: 10, tibia: 5, tarsus: 9.

Scutum: Measurements (1 specimen); ASL: 104, SB: 40, TS: 46,
ASL/SB: 2.6.

Remarks. — The key couplet separating T. kansasensis from T.
gurneyi uses the longer cheliceral base of kansasensis. A second
character is the length of the posterior body setae; those of kansas-
ensis are twice as long as those of gurneyi.

Specimens examined. — Total 1 nymph, as follows: Kansas. Bar-
ber County: 4 mi. S Aetna, ISJeotoma micropus, July 11, 1955 (1
nymph, reared DAC, idet. by larval skin).

Tromhicula kardosi Loomis
(Figs. 59, 78, 161, 174, 175)

Tromhicula kardosi Loomis, 1954, Univ. Kansas Sci. Bull., vol. 36, pp. 929-930

( larva ) .
Tromhicula (Neotromhicula) kardosi, Loomis, 1956, Univ. Kansas Sci. Bull.,

vol. 37, p. 1299 (larva).

Description of nymph. — Resembles T. ftchi except as follows:

Body: About 0.6 mm. long, color orange. Setae similar to those
of T. fitchi, but posterior setae with occasional long branches among
shorter branches of distal halves; posterior setae about 65 pi long.
Measurements of tarsus I (means of 7 specimens); TL: 108 ± 1.091,
TH: 56.1 ± 1.534, TL/TH: 1.93 ± 0.0565.

Gnathosoma: Cheliceral blade with about 17 small teeth. Chelic-
eral measurements (means of 7 specimens); BL: 111.0 ±: 0.7228,
BH: 38.6 ± 1.042, CL: 54.4 ± 0.8401, BL/BH: 2.89 ± 0.0799,
BL/CL: 2.04 ± 0.0368. Palpus with tarsala about 11 jjl long. Setal
counts; femur (5 specimens): 3.3 ± 0.1225; genu (6 specimens):
7.83 ± 0.1923, tibia: range 3-5, tarsus: range 8-9.

Scutum. Generally resembles that of T. fitchi; sensilla usually
with more numerous branches, resembling those of T. lipovskyi.
Scutal measurements (means of 6 specimens); ASL: 94.2 ± 2.057,
SB: 36.8 ± 0.4474, TS: 27.6 ± 0.2974, SENS: about 125, ASL/SB:
2.57 ± 0.0761.

Specimens examined. — Total 7 nymphs, as follows: Kansas.
Douglas County: 4/2 mi. W, 3 mi. S Baldwin, Sciurus niger, No-
vember 28, 1951 (7 nymphs, reared DAC).

264 The University Science Bulletin

Trombicula merrihewi Loomis and Lipovsky

(Figs. 38, 89, 136, 137, 165, 181, 232)

Trombicula merriheici Loomis and Lipovsky, 1954, Jour. Kansas Ent. Soc,
vol. 27, p. 51 (larva).

Description of nymph. — Resembles T. crossleyi except as follows:

Body: About 0.6 mm. long, color not known. Body setae short;
branches fine, long, rather sparse; tips of posterior setae of 2 or 3
branches; posterior setae about 14 \i long. Measurements of tarsus
I (means of 10 specimens); TL: 69.0 ± 0.6146, TH: 36.5 ± 0.4281,
TL/TH: 1.89 ± 0.0278.

Gnathosoma: Cheliceral base punctate; blade with about 14
small teeth. Cheliceral measurements (means of 10 specimens);
BL: 76.8 ± 0.827, BH: 28.2 ± 0.512, CL: 33.0 ± 0.516, BL/BH:
2.75 ± 0.0401, BL/CL: 2.32 ±: 0.0359. Palpus with claw more
normal in shape; tibial setae nude or nearly so; tarsus with setae
with reduced branching, tarsala about 7 \). long. Setal counts
(means of 10 specimens); femur: 3.0 (no variation); genu: 4.75 ±:
0.1539, tibia: range 4-5, tarsus: range 8-9.

Scutum: Tectum with about 8 prominent teeth on margin. Sen-
sillary area triangular in appearance; strong ridges posterior to
sensillary bases, others weak; bulla apparently unusually elevated;
carina thick; without unsually elongated parascutal setae; sensilla
subclavate. Scutal measurements (means of 10 specimens); ASL:
59.1 ± 1.120, SB: 25.1 ± 0.888, TS: 19.4 ± 0.612, SENS: about 60,
ASL/SB: 2.38 ±0.0789.

Remarks. — This species is readily recognized by the unusually
short (14 \).) posterior body setae.

The occurrence of larvae on bats strongly suggests that the
nymphs inhabit caves. Of the characters given above, the reduced
branching of the palpal setae seems most likely to be a modification
for this habitat. Of the species probably dwelling in caves as
postlarvae, Speleocola tadaridae also has reduced branching of
palpal setae, but Whartonia sensae and Euschdngastia pipistrelli do
not. Also, Trombictda ornata, a species related to T. merrihewi,
but not taken from bats, shows a similar reduction in branching.

Specimens examined. — Total 19 nymphs, as follows: Oklahoma.
Woods County: 6 mi. S, 2 mi. W Aetna, Kans., Tadarida mexicana,
August 24, 1949 (19 nymphs, reared LJL).

Nymphs of the Trombiculidae 265

Tromhicula montanensis Brennan

(Figs. 9, 12, 54, 73, 122, 123, 167, 239)

Tromhicula montanensis Brennan, 1946, Jour. Parasit., vol. 32, pp. 441-442
(larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1306-1310 (larva).

Description of nymph. — Body: About 0.6 mm. long, rounded,
color yellow to orange. Setae with thick stems, rather short;
branches fine, numerous; posterior setae with tips somewhat
rounded, with several tiny branches; posterior setae about 30 [x
long. Sternal area variable; usually closed behind but rarely open;
usually about pentagonal in shape but rarely elongated-rectangular;
with about 8 setae. Measurements of tarsus I (means of 17 speci-
mens); TL: 89.9 ±1.6997, TH: 45.1 ± 0.8613, TL/TH: 1.99
± 0.0251.

Gnathosoma: Cheliceral base of usual shape, punctate; blade nor-
mal, with about 13 teeth of intermediate size. Cheliceral measure-
ments (means of 9 specimens); BL: 90.8 ±: 1.9985, BH: 33.6 ±
0.7094, CL: 46.2 ± 0.8127, BL/BH: 2.71 ±: 0.0611, BL/CL: 1.96
± 0.0176. Palpus of usual shape; dorsal genual setae often nude;
claw and accessory claws rather small; dorsal tibial setae oc-
casionally nude; tarsus with tarsala about 9 [jl long; tarsus with
3 inner, 1 outer apical nude setae. Setal counts (means of 9 speci-
mens); femur: 4.00 ± 0.2079, genu: 8.28 rh 0.1900, tibia: range
4-5, tarsus: 9.

Scutum: Tectal margin with about 10 teeth. Sensillary area
triangular in appearance; basal plate rather simple, of two halves;
apodeme well developed; bulla triangular, prominent, conspicuously
domelike; carina thick, only slightly elevated, often difficult to
distinguish from bulla; posterior ridge weak, ridges medial to sen-
sillary bases weak, others strong; sensilla with short branches basally
but longer ones distributed over rest of stems, stems flexible, flagel-
liform or slightly thickened. Scutal measurements (means of 16
specimens); ASL: 71.8 ± 0.9593, SB: 32.4 ± 0.7161, TS: 22.7
± 0.506, SENS: about 80, ASL/SB: 2.26 ± 0.0418.

Remarks. — Tromhicula montanensis is a common chigger of the
central states region. The best recognition character for nymphs
seems to be the four apical nude setae of the palpal tarsus; other
species of Tromhicula (Tromhicula) have either three, five, or six
such setae.

266 The University Science Bulletin

Nymphs of T. montancnsis show considerable variation in mor-
phology. The shape of the sternal area, which is relatively con-
stant for other species, is dimorphic in montanensis; occasional
specimens from Barber County, Kansas, possess an elongated-
rectangular sternum which is open behind, instead of the usual
closed pentagonal sternum. Other variation is noticeable in the
scutal area. A carina is plainly visible on some specimens but is
not visible on other equally well-mounted specimens. A dimorphic
condition of the sensilla occurs also; the two distinct forms are a
flagelliform stem resembling that of T. gitrneyi and a slightly thick-
ened stem resembling that of T. Upovskyi. The more common type
is the latter one, but occasional specimens from Barber County,
Kansas, and Webster County, Nebraska, have flagelliform sensilla.
The flagelliform type is more prevalent in a series of 68 nymphs
from Barber County collected in August of 1948, but is uncommon
on specimens taken in 1949, 1952, and 1953 in Barber County.
Associated larvae show no unusual variation.

The reasons behind these unusual variations are not known. The
specimens at hand are not sufficient to indicate whether genetic
or environmental forces produce these eftects.

Nymphs and adults of T. montanensis feed readily upon collem-
bolan eggs. This species has been reared through a complete
generation.

Specimens examined. — Total 108 nymphs, as follows: Kansas.
Barber County: 5 mi. E Aetna, Ferognathus hispidus, July 26, 1952
(4 nymphs, reared DAC, 1 idet. by larval skin); 43-2 mi. S, 1 mi. W
Aetna, Citellns tridecemlineatiis, September 16, 1953 (2 nymphs,
reared DAC) and Cynomys hidovicianus, September 16, 1953 (1
nymph, reared DAC); 4 mi. S Aetna, Neotoma micropus, August 22,
1948 (68 nymphs, reared LJL); 10/2 mi. W Hardtner, Cynomys
hidovicianus, August 21, 1949 (5 nymphs, reared LJL), and July
26, 1952 (7 nymphs, reared DAC); 3/2 mi. W Hardtner, Cynomys
hidovicianus, August 23, 1949 (1 nymph, reared LJL). Rawhns
County: 11 mi. S McDonald, Cynomys hidovicianus, July 27, 1948
(1 nymph, reared LJL). Seward County: 4 mi. NE Liberal, Mas-
ticophis flagelhim, September 12, 1948 (1 nymph, reared LJL).
Nebraska. Hitchcock County: 4 mi. E Stratton, Cynomys ludo-
vicianus, August 8, 1949 ( 1 nymph, reared LJL ) ; 13 mi. S Trenton,
CiteUus tridecemhneatus, August 8, 1949 (9 nymphs, reared LJL).

Nymphs of the Trombiculidae 267

Webster County: 3 mi. E Guide Rock, Heterodon nasicus, August 5,
1951 (7 nymphs, reared D AC). Oklahoma. Woods County: 6 mi.
S, 2 mi. W Aetna, Kans., Heterodon platyrhinos, October 7, 1951
( 1 nymph, reared DAC ) .

Trombicula ornata Loomis and Lipovsky

(Figs. 58, 162, 202)

Trombicula ornata Loomis and Lipovsky, 1954, Jour. Kansas Ent. Soc, vol. 27,
p. 47 {larva).

Description of nymph. — Resembles T. crossleyi except as follows:

Body: About 0.7 mm. long, rather rounded, color not known.
Posterior setae about 25 ]}. long. Sternal area larger than that of
T. crossleyi, with about 4 to 6 setae. Measurements of tarsus I
(means of 7 specimens); TL: 90.1 ± 2.176, TH: 38.7 d= 0.6801,
TL/TH: 2.34 ± 0.0812.

Gnathosoma: Cheliceral base unusually elongated, area of articu-
lation elongated; blade rather stout but shearlike, with about 17
small teeth. Cheliceral measurements (means of 7 specimens);
BL: 111.3 ± 1.229, BH: 30.7 ± 0.5217, CL: 39.4 ± 0.7189, BL/BH:
3.63 ± 0.1286, BL/CL: 2.83 ± 0.0474. Palpus with claw more nor-
mal in size; accessory claw thin; tibial setae nude or nearly so;
tarsala about 9 ^ long. Setal counts (6 specimens); femur: 3.00
(no variation); genu: 4.79 zt 0.4062, tibia: range 4-5, tarsus: 9.

Scutum: Sensillary area markedly triangular in shape; strong
ridges anterior to and posterior to sensillary bases, others weak;
sensillary bases not unusually prominent. Scutal measurements
(means of 6 specimens); ASL: 72.2 ± 0.9700, SB: 28.9 + 0.5084,
TS: 24.7 ± 0.7603, SENS: about 75, ASL/SB: 2.48 ± 0.0538.

Remarks. — This species appears most similar to Trombicula
merrihewi, although similarities are more obvious in the larval stage
than in the nymphal stage. Trombicula ornata resembles the larger
Trombicula species almost as much as it resembles T. crossleyi and
relatives, but the affinities of ornata] through merriheivi, would seem
to be with crossleyi and trisetica.

Trombicula ornata is easily separated from these species by the
elongated cheliceral base which it possesses.

Specimens examined. — Total 18 nymphs, as follows: Kansas.
Barber County: 3^2 mi. S, 1 mi. W Aetna, Neotoma micropus,
April 11, 1949 (16 nymphs, reared LJL); 4 mi. S Aetna, Neotoma
micropus, August 22, 1949 (2 nymphs, reared LJL).

268 The University Science Bulletin

Trombicula trisetica Loomis and Crossley

(Figs. 1-7, 198)

Trombicula trisetica Loomis and Crossley, 1953, Joiir. Kansas Ent. Soc., vol. 26,
p. 32 (larva); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1321-1323
(larva).

Description of nymph. — Resembles T. crossley i except as follows:

Body: About 0.6 mm. long. Posterior setae about 25 [x long.
Measurements of tarsus I (means of 10 specimens); TL: 82.8 =h
1.397, TH: 40.1 ±: 0.647, TL/TH: 2.09 ± 0.0277.

Gnathosoma: Cheliceral base punctate; blade with about 17 small
teeth. Cheliceral measurements (means of 10 specimens); BL:
90.5 ± 1.046, BH: 30.4 ± 0.3711, CL: 40.5 ± 0.2687, BL/BH;
2.91 ± 0.0407, BL/CL: 2.19 ± 0.0278. Palpus with tarsala about
9 [i long. Setal counts (means of 10 specimens); femur: 2.95 ±
0.0049, genu: 3.00 (no variation); tibia: range 5-6, tarsus: 9.

Scutum: Sensilla less expanded than those of T. crossJeyi, distal
branches somewhat longer. Scutal measurements (means of 10
specimens); ASL: 66.1 ± 1.059, SB: 30.3 ± 0.7753, TS: 22.8 dz
0.4163, SENS: about 85, ASL/SB: 2.21 ± 0.0722.

Specimens examined. — Total 18 nymphs, as follows: Kansas.
Douglas County: Univ. Kansas Natural History Reservation, Elaphe
ohsoleta, September 10, 1952 (18 nymphs, reared DAC).

Genus Walchia Ewing

Walchia Evdng, 1931, Proc. U. S. Nat. Mus., vol. 80, p. 10, [type Trombidium
glabrum Walch (not Trombidium glabrum Duges) := Walchia pinquf,
Gater] (larva); Wharton et al, 1951, Jour. Parasit., vol. 37, p. 30 (adult);
Wharton and Fuller, 1952, Mem. Ent. Soc. Washington, no. 4, p. 91
(postlarvae); Audy, 1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 161
(adult); Loomis, 1956, Univ. Kansas Sci. Bull., vol. 37, p. 1362 (larva).

Gahrliepia (Walchia), Womersley, 1952, Rec. South Australian Mus., vol. 10,
p. 393 (nymph).

Diagnosis (nymph). — (1) Eyes absent. (2) Precoxal plates of
coxae I absent. (3) Body setae branched, not expanded, without
compound branching; adjacent setae of similar lengths. (4) Claws
on tarsus I undivided distally. (5) Tarsus I with dorsoapical stump-
like process. (6) Cheliceral base not unusually elongated; blade
shearlike. (7) Palpal trochanter not elongated. (8) Hypostome
short, blunt; with about twenty short apical nude or nearly nude
setae. (9) Pharynx marked with a reticulate pattern. (10) Tectum
almost square, hyahne, margin with teeth; ivithout tectal seta. ( 11 )
Sensillary area wide; mean ratio ASL/SB: 1.3. (12) Without
branched body seta in sensillary area. (13) Sensilla straight, with
branches.

Nymphs of the Trombiculidae 269

Additional features (nymph). — (14) Body small (about 0.5 mm.
long). (15) Sternum elongated-rectangular in shape, closed be-
hind. (16) Tarsus I elongated. (17) Artictdating area of cheliceral
base on posterior margin; posterior tip of cheliceral base elevated.

(18) Palpus stout, small; claw slightly elongated, rather straight;
distal portion of tibia not unusually elongated; two accessory claws;
tarsus with seven to nine branched setae and one apical nude seta.

( 19) Hypostome with about ten ventral branched setae. (20) Basis
capituli of normal shape. (21) Crista a short narrow rod. (22)
Sensillary area with basal plate of two well-separated halves; bulla
rather large; carina absent; sensillary bases placed well lateral; an-
terior and posterior ridges present; lateral ridges present but less
prominent than ridges medial to sensillary bases; apodeme small.
( 2-3 ) Only two parasctital setae.

Remarks. — One species of the genus, Walchia americana, is known
from the United States. Ten or more species are known from the
Asiatic-Pacific Begion. Nymphs of four of these species have been
described by Womersley (1952:285).

Womersley (1952:278) considered Walchia, Schongastiella Hirst,
and Gateria Ewing to be subgenera of Gahrliepia Oudemans. This
opinion was based upon comparisons of both larvae and nymphs of
members of these genera. Womersley 's opinion may certainly be
correct; the features mentioned in his descriptions of nymphs do
not appear to indicate generic separation for these forms. I am
persisting, however, in using the generic name Walchia (as did
Loomis, 1956), at least until Womersley's more conservative ap-
proach can be substantiated.

While Womersley's descriptions are not adequate for close com-
parisons, it does appear that at least some of the species he describes
will not key out to Walcliia in my key. In the generic diagnoses for
Gahrliepia, Womersley states that ,one epistomal seta ( tectal seta )
is present. However, this seta was clearly seen by him on only two
of ten species, and was but indistinctly seen on two others. The
tectal seta is definitely absent on both nymphs and adults of
Walchia americana, and I have included this as a generic feature
and used it in the key to genera, to separate Walchia from
Schoiitedenichia. If Womersley's descriptions are correct some
additional feature must be found. The single pair of parascutal
setae or the single apical nude seta of the palpal tarsus of Walchia
possibly separate these two genera; unfortunately these features
are not mentioned by Womersley.

270 The University Science Buixetin

Walchia americana Ewing

(Figs. 35, 84, 118, 119, 147, 188, 203, 235)

Walchia americana Ewing, 1942, Jour. Parasit., vol. 28, p. 491 (larva); Loomis,
1956, Univ. Kansas Sci. Bull., vol. 37, pp. 1363-1365 (larva).

Descriptioji of nymph. — Body: About 0.5 mm. long, rounded,
color white. Setae short, stems thick at bases and abruptly tapered,
with many short branches; posterior setae with tips of 2-3 tiny
branches; posterior setae about 14 i). long. Sternum closed behind,
elongated posteriorly, often with an indication of incipient longi-
tudinal division; with about 8 setae. Tarsus I with small stumplike
process projecting from apical dorsal margin. Measurements of
tarsus I (means of 7 specimens); TL: 77.9 zt 1.945, TH: 46.7 +
1.646, TL/TH: 1.67 ± 0.0606.

Gnathosoma: Cheliceral base of characteristic shape, punctate;
blade shearlike, with about 10 small teeth. Cheliceral measurements
(means of 10 specimens); BL: 56.3 ± 0.731, BH: 28.0 =t 0.4899,
CL: 28.0 ± 0.648, BL/BH: 2.01 ± 0.0434, BL/CL: 2.01 ± 0.0456.
Palpus with accessory claws long; tarsus with tarsala about 9 ;ji long,
apical nude seta on inner surface. Setal counts ( means of 10 speci-
mens); femur: 4.95 ± 0.0879, genu: 7.4 ± 0.2082, tibia: range 3-4,
tarsus: range 7-9.

Scutum: Sensillary area broadly transverse-oval; apodeme re-
duced; major posterior ridge rrmning medial to sensillary bases;
ridges anterior and posterior to sensillary bases weak; posterior
ridge discontinuous, ends joined by small secondary ridge; bulla
triangular, prominent; carina absent; sensilla with stems thickened
but flexible, with short basal branches and longer branches on re-
mainder of stems. Scutal measurements (means of 10 specimens);
ASL: 45.1 ± 0.836, SB: 34.6 ± 0.7023, SENS: about 75, ASL/SB:
1.32 ± 0.0326.

Remarks. — In Womersley's (1952:280) key to species of Gahrlie-
pia, Walchia americana will key out to G. rtistica, from which it may
be separated by the shorter posterior body setae (14 \). long in
americana, 32 [i long in rtistica).

Walchia americana was reared on four occasions and reached a
complete generation on three of these occasions. The nymphs and
adults were fed upon active stages of the collembolan, Sinellu
curviseta.

Specimens examined. — Total 21 nymphs, as follows: Kansas.
Douglas County: Sciurus niger, January 19, 1950 ( 2 nymphs, reared
LJL); 2 mi. S Warden, Sciurus niger, November 28, 1949 (1 nymph.

Nymphs of the Trombiculidae 271

reared LJL ) ; 4/2 mi. W, 3 mi. S Baldwin, Sciurus niger, November 28,
1951 (4 nymphs, reared DAC). Jefferson County: 10 mi. W Mid-
land, November 14, 1953 (14 nymphs, reared RBL-DAC).

Additional Genera Known As Postlarvae

This section includes those genera which are known as post-
larvae but for which no specimens were available for study. Most
are included in the key to genera on characters given in descriptions
by other authors. Many of these descriptions are brief; often they
contain no indication of characters considered as generic in the
present paper. Thus it is not possible to give full diagnoses for
these genera. In the discussions, further features are indicated,
usually by comparisons with better known genera.

Subfamily Trombiculinae Ewing
Genus Ascoschongastia Ewing

Ascoschongastia Ewing, 1946, Proc. Biol. Soc. Washington, vol. 59, p. 71 (type

Neoschiingastia malaijensis Gater) (larva).
Schongastia (Ascoschongastia) , Womersley, 1952, Rec. South Australian Mus.,

vol. 10, p. 384 (part.) (nymph).

Diagnostic features (nymph). — Apparently identical with Eu-
schongastia.

Remarks. — This genus is recognized by Wharton and Fuller
(1952:71) for trombiculine species which as larvae have expanded
sensilla and the PL setae oflF of the scutum. Five species are included.
Womersley (1952:384) described the nymph of A. malayensis and
concluded that it was not generically separable from those of species
which he placed in Schongastia (Ascoschongastia) (:= Euschon-
gastia). Womersley confirms in a footnote (ihid, p. 170) that
Euschongastia has priority over Ascoschongastia and that this
synonomy should obtain in his system.

The description of the nymph of A. malayensis as given by
Womersley is not adequate for purposes of comparison. Therefore,
I must follow Womersley in regarding malayensis as inseparable
from nymphs of Euschongastia.

Genus DoJoisia Oudemans

Doloisia Oudemans, 1910, Ent. Bcr. Amst., vol. 3, p. 87 (type Doloisia sijnoti
Oudemans) (larva); Sasa, 1953, Japanese Jour. Exp. Med., vol. 23, p. 427
(ntjmph); Audy, 1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 1.57
(nijriiph).

Diagnostic features (nymph). — (1) Eyes absent. (2) Precoxal

plates absent. (8) Hypostome pointed, projecting, with about

eight apical nude setae. (10) Tectum with teeth on margin. (11)

Sensillary area broad, ratio ASL/SB: 0.7-1.2

272 The Uni\'ersity Science Bulletin

Remarks. — The nymph of Doloisia okahei Sasa et ah, has been
well described by Sasa (1953:427); the nymph of D. ocuUcola
(Womersley) has been described by Womersley (1952:383); also,
Audy (1954:157) has given some nymphal characters. The above
diagnosis is based upon these publications. From Sasa's figures it
appears that the palpal shape and the apical nude setae of the
hypostome may be distinctive, as well.

Larvae of Doloisia have small characteristic scuta with expanded
sensilla.

The general facies of nymphs as presented by authors is distinc-
tive; the scutum of the nymph is somewhat similar to that of Gun-
therana but the species are abundantly distinct from that genus.

Genus Gahrliepia Oudemans

Typhlothrombium Oudemans, 1910, Ent. Ber. Amst., vol. 3, p. 105 (type
Typhlothrombium nanus Oudemans) (larva) (not Typhlothrombium
Berlese).

Gahrliepia Oudemans, 1912, Ent. Ber. Amst., vol. 3, p. 273 (type Typhlo-
thrombium nanus Oudemans) (larva); Womersley, 1952, Rec. South
Australian Mus., vol. 10, p. 393 (nymph).

Gateria Ewing, 1938, Jour. Washington Acad. Sci., vol. 28, p. 295 (type
Gahrliepia fietcheri Gater) (larva).

Schongastiella Hirst, 1915, Bull. Ent. Res., vol. 6, p. 188 (type Schongastiella
bengalensis Hirst) (larva).

Diagnostic features {nymph). — Same as those for Walchia.

Remarks. — Womersley (1952:279) synonomized the genera Ga-
teria, Schongastiella, and Walchia with Gahrliepia, on both larval
and postlarval evidence. Some authors have been somewhat re-
luctant to accept this synonomy (see Audy 1954:161). Womersley
described nymphs for species in all four genera, and on the basis of
his descriptions, they are only specifically separable. I have used
the generic name Walchia for W. americana and based my diagnosis
of the nymph on that species. It is difficult to say how closely
Womersley 's species fit that diagnosis (see discussion of Walchia).
Thus it is impossible for me to evaluate the Gahrliepia complex.
Quite possibly Womersley 's solution is the correct one.

Genus Gtmtherana Womersley and Heaslip

Guntherana Womersley and Heaslip, 1943, Trans. Roy. Soc. South Australia,
vol. 67, p. 132 (type Neoschongastia kallipygos Gunther) (larva); Wharton
and Fuller, 1952, Mem. Ent. Soc. Washington, no. 4, p. 83 (larva, post-
larva); Womersley, 1952, Rec. South Australian Mus., vol. 10, p. 372
(postlarva); Audy, 1954, Stud. Inst. Med. Res. Fed. Malaya, no. 26, p. 156
(larva, postlarva).

Diagnostic features {nymph). — (2) Precoxal plates of coxae I
present but not fused in midline to form longitudinally divided

Nymphs of the Trombiculidae 273

sternum. (3) Body setae branched; adjacent setae of similar lengths.
(11) Sensillary area wide, ratio ASL/SB: about 1.1.

Remarks. — These features are taken from Womersley (1952:372)
and Audy (1954:156). Presumably they hold for the nymph and
adult of G. kalUpygos, and for G. tindalei and G. translucens, the
latter known only as adults. Larvae of Gimtherana are distinguished
by the presence of caudal plates.

Relationships of the genus Guntherana are hard to define on the
basis of the brief published descriptions. Nymphs appear to re-
semble those of Trombicula; however, the presence of precoxal
plates indicates similarity with Neoschongastia and the scutum re-
sembles that of members of the Eiisclidngastoides-Pseudoschdn-
gastia-Walchia group of genera. A detailed study of nymphs of
the genus Guntherana would be of the greatest interest.

Genus Heaslipia Ewing

Tromhiculoides Womersley and Heaslip, 1943, Trans. Roy. Soc. South Aus-
tralia, vol. 67, p. 101 (type Tromhiculoides gateri Womersley and Heaslip)
(larva) (not Tromhiculoides Jacot).

Heaslipia Ewing, 1944, Proc. Biol. Soc. Washington, vol. 57, p. 103 (type
Tromhiculoides gateri Womersley and Heaslip) (larva); Womersley, 1952,
Rec. South Austrahan Mus., vol. 10, p. 422 (mjmph); Audy, 1954, Stud.
Inst. Med. Res. Fed. Malaya, no. 26, p. 149 (nymph).

Diagnostic features (nyniph). — (10) Tectum large, an elongate
triangular cone, not hyaline, margin without teeth or serrations.
Otherwise similar to Eiischongastia.

Remarks. — This diagnosis is after Womersley (1952:422) and

Audy (1954:149) based upon Heaslipia gateri. Quite possibly the

conelike tectum will not distinguish nymphs of Heaslipia from

nymphs of some Eiischongastia. Larvae are similar to those of

Trombicula but have additional scutal setae. The genus is mono-

typic.

Genus Ipotrombicula Womersley

Ipotromhicula Womersley, 1952, Rec. South Australian Mus., vol. 10, p. 323
(type Tromhicula elegans Womersley) (adult only).

Diagnostic feature (adult). — (12) With branched body seta in
sensillary area.

Remarks. — This genus is monotypic for I. elegans, known only as
adult. Except for the key character of the possession of a branched
seta in the sensillary area, this species would probably be placed
in Trombicula, subgenus Eutrombicula; it possesses eyes in typical
Eutrombicula position. Indeed this may be its proper place.

274 The University Science Bulletin

Genus Schongastia Oudemans

Schdngastia Oudemans, 1910, Ent. Ber. Amst., vol. 3, p. 87 (type Thrombidium
vandersandei Oudemans) (larva); Womersley, 1952, Rec. South Australian
Mus., vol. 10, p. 376 (nymph); Audy, 1954, Stud. Inst. Med. Res. Fed.
Malaya, no. 26, p. 150 (nymph).

Diagnostic features {nymph). — (2) Precoxal plates of coxae I
present but not fused in midline to form longitudinally divided
sternum. (3) Adjacent body setae of similar lengths. (11) Sen-
sillary area triangular; ratio ASL/SB: about 2.0.

Remarks. — Nymphs of three species of Schdngastia were de-
scribed by Womersley (1952:381-387). Womersley does not men-
tion the precoxal plates, but Audy (1954:150) says that these are
present. The nymphs, as described by Womersley, show no other
features clearly separating them from Trombicula.

Genus Schoiitedenichia Jadin and Vercammen-Grandjean

Schoutedenichia Jadin and Vercammen-Grandjean, 1954, Ann. Mus. Congo
Tervuren, Zool., vol. 1, p. 195 (type Schoutedenichia ftdleri Jadin and
Vercammen-Grandjean) (larva).

Diagnostic features {nijniph). — (2) Precoxal plates of coxae I
absent. (5) Tarsus I with dorsoapical stumplike process. (10)
Tectum with a single tectal seta.

Remarks. — These features are taken from descriptions by Jadin
and Vercammen-Grandjean (1954:287, 289) of nymphs of S. pene-
trans and S. pirloti. In combination these features separate nymphs
of Schoutedenichia from those of other genera. From the illustra-
tions by Jadin and Vercammen-Grandjean it may be seen that the
genus has some features of the Euschdngastoides-Pseudoschdn2.astia-
Walcliin group of genera, including the scutal shape, the blunt
hypostome with about twenty short apical setae, and the features
of palpal setation. As previously noted, Schoutedenichia is very
probably closely related to this group of genera.

Genus Speotrombicula Ewing

Speotrombicula Evidng, 1946, Jour. Parasit., vol. 32, p. 437 (type Tromhicula
trifurca Evi^ing) (adult only); Wharton and Fuller, Mem. Ent. Soc. Wash-
ington, no. 4, p. 90 (adult); Womersley, 1952, Rec. South Australian Mus.,
vol. 10, p. 324 (adult).

Diagnostic feature (adult). — (4) Claws on tarsus I trifurcate
distally.

Remarks. — Speotrombicula is monotypic for S. trifurca, known
only as adult. Judging from E wing's (1933:3) description the
species is a Trombicula except for the key character of the divided
claws. However, this description is brief.

Nymphs of the Trombiculidae 275

Genus Walchiella Fullea-

Walchiella Fuller in Wharton and Fuller, 1952, Mem. Ent. Soc. Washington,
no. 4, p. 95 (type Tromhicula oudemansi Walch) (larva).

Diagnostic features (nymph). — None.

Remarks. — Womersley (1952:380) has described the nymph of
Walchiella oudemansi, which he inckided in Schongastia (Schon-
gastia). His description indicates no features which separate the
nymph from those of Tromhicula. A possible useful character is
present in the posterior body setae, which have a subterminal
branch that is unusually long (as long as the seta itself). Consider-
ing the variation in form of the posterior body setae, as observed in
Euschongastia and Tromhicula, this character does not seem to
have generic significance.

Walchiella is monotypic for W. oudemansi. Larvae have six seg-
ments in legs II and III, expanded sensilla, and five scutal setae.

COMPARISON OF SYSTEMS OF CLASSIFICATION

Workers are in general agreement concerning the classification
based primarily on larvae. In the main similar generic levels are
given in recent papers, but there is less agreement on generic align-
ment in subfamilies. Andy (1954:134-135) presents in tabular form
his own generic arrangement, that of Wliarton and Fuller (1952),
and that of Womersle>' ( 1952 ) . Although these systems are di-
vergent it may be seen that there is general agreement as to the
status of most of the genera.

These systems of classification are based primarily on the larval
stage but taxonomic characters of known postlarvae were quite
evidently considered. The classification in the present paper is
based upon both larval and postlarval stages, perhaps with the
latter stages given more than the usual weight.

Concerning the species level, there appears to be no reason for
insisting that species be distinct from one another in both larval
and postlarval stages. The majority of the species studied are dis-
tinct in both stages, although some ( for example, species of Pseudo-
schongastia) are more readily separated as larvae while others
(species of Neoschongastia) are more readily separated as nymphs.
An area of greater interest in the comparison of systems is at the
more arbitrary generic and higher levels, where weighting of
characters becomes important.

With the two classifications there are four types of generic cate-
gories as regards distinctiveness; these are (1) genera distinct, /. e..

276 The Univ^ersity Science Bulletin

recognizable by a character or combination of characters, in both
the larval and postlarval stages, (2) genera distinct in the larval stage
but not recognizable in postlarval stages, (3) genera distinct in
postlarval stages but not recognizable in the larval stage, and (4)
genera distinct in neither stage but recognizable by some other
criterion. Examples of the first three of these points are known.
These considerations, of course, apply to subgenera, subfamilies,
and other supraspecific categories as well as to genera.

Most of the subgenera and genera were found to be recognizable
on morphological characters of both larval and postlarval stages.
Members of the subgenus Neotrombictila are recognizable on larval
characters but no postlarval characters were found for this group.
Species in the genus Bhnkaartia and perhaps the subgenus Lepto-
trombidhim are diflBcult to recognize as larvae but are readily recog-
nized on postlarval characters. The genus Etischongastoides, as
recognized in the present paper, contains two species clearly distinct
as nymphs but as larvae they are members of different genera.
With these few exceptions, evidence of the nymphal stage corrob-
orates the classification based on larvae.

Of the exceptions noted above, Blankaartia and Neotrombictila
are not serious discrepancies; in both cases relationships are clear
in larval and postlarval stages. In other genera, also, the degree
of distinctiveness is not the same in the two phases of the life
history. Some as yet undetected character may serve to separate
nymphs of species of Neotrombictila from those of the typical sub-
genus. Further studies of the larvae of Blankaartia may make the
identification of larvae of that genus easier. In these cases evidence
from postlarval stages does not show that relationships are other
than those indicated in the larval stage; the matter is one of relative
distinctiveness.

A question posed by these exceptional cases concerns the status
of genera and subgenera; namely, should a genus (or subgenus)
necessarily be recognizable in both larval and postlarval stages in
order to be acceptable? Womersley, in a revision of trombiculids,
considered postlarval evidence and concluded (1952:9), "Such a
revision shows that many of the larval genera which have been pro-
posed cannot be separated in the nymphal or adult stages, should
not be used except perhaps for convenience, as subgenera based on
larval features only." Audy, in consideration of this problem writes
(1954:127), ". . . rigid ideas on the employment of either
larval or adult characteristics are to be deplored." "Certainly,

Nymphs of the Trombiculidae 277

species and subgenera based entirely on morphological larval
characters may be fully acceptable though the free extension of
this to genera (and even to subfamilies, as has been done) is de-
batable in particular instances." The present writer is more in
agreement with Audy than with Womersley. It does not seem pos-
sible to draw a hard and fast rule for cases of relative distinctiveness.
The creation of genera indistinguishable in one phase of the life
history poses practical problems rather than philosophical ones.

The genus Euschongastoides poses a more difficult problem. As
explained in the taxonomic section of this paper, the two species
of Euschongastoides, as larvae, key out to different genera, Trom-
bicula and Eiischongastia. Except for the key character, the nature
of the sensillum, the two larvae are very similar. As nymphs, the
two species are similar to each other but differ markedly from
Trombictda and Euschongastia. The relationships of the two species
(on nymphal evidence) appear to be with Pseudoschongastia and
Walchia. If the relationships as indicated by the nymphs are cor-
rect, then the erroneous placement of the larvae was due to the
rigid use of a single key character in separating Trombictda and
Euschongastia. If the relationships are correct on larval evidence,
the similarity of the nymphs ( and of other larval characters ) must
be assumed to be due to parallelism. The writer feels that the
nymphal evidence is the more convincing and thus gives generic
statvis to Euschongastoides.

Undoubtedly other similar cases will arise. If these are judged
on their individual merits, without undue weighting of characters
of any stage, the result should be a more natural system of classifica-
tion. The postlarval evidence should be considered as additional
knowledge concerning the relationships of the organisms.

The remaining genera are distinct as such in both phases of the
life history. Both Euschongastia and Speleocola seem closely re-
lated to Trombicida in both stages.' In Cheladonta and Neoschon-
gastia, the larvae are similar to those of Euschongastia, but post-
larvae of these two genera are quite distinctive. Pseudoschongastia
and Walchia are similar to one another as nymphs, more so than
as larvae.

RUDIMENTS OF PHYLOGENY

As previously stated, this study is restricted mainly to species in
North America, and the available nymphs represent but a fraction
of the species known from this continent. Some genera are known
only as larvae. Under these conditions it would not be realistic to
attempt to construct a detailed phylogeny of the group. However,

278 The University Science Bulletin

it seems possible that an outline of the evolutionary development of
the group might be proposed, since at least the major genera of
trombiculids are known as nymphs. The following ideas, therefore,
are oflFered as the writer's opinions; the evidence for them is meager
in most cases and further work will undoubtedly indicate modifica-
tions. The genera not studied by the writer are not included in this
discussion; to do so would be compounding speculation.

The origin of trombiculids is almost certainly in the family Trom-
bidiidae. The distinction between the two families is not clear-cut.
Certain genera, such as Ncotwmbidiuw, could be placed in either
family on morphological grounds (Borland, 1956:30). From evi-
dence available at present, it appears that trombiculids are closely
related to members of the trombidiid subfamily Microtrombidiinae,
which itself is heterogeneous. Trombiculids are not far removed
from the primitive trombidiids (Johnstoniana and relatives).

By analogy with the trombidiids, certain characters of some trom-
biculid nymphs are deduced to be primitive. The primitive condi-
tion of the gnathosoma appears to be as follows: Hypostome
pointed, projecting, with about eight apical nude setae. Cheliceral
base elongated, with posterior tip depressed; cheliceral blade shear-
like, with teeth. Palpus large, with numerous setae, with claw and
two accessory claws, and tarsus with several apical nude setae.
Primitive features of the scutum include: Tectum rounded, margin
smooth; with tsvo tectal setae. Crista rodlike, long. Sensillary
area simple, circular or oval shaped, with few or no ridges; sensilla
without branches; posterior apodeme rodlike. Other primitive
characters appear to be the presence of eyes anterior to the sen-
sillary area, the absence of a closed sternum, and a simple circular
anus. While unmodified body setae are possibly primitive, so many
genera in the family Trombidiidae have highly modified body setae
that it seems unsafe to make this generalization for trombiculids.
The strongly constricted body of most trombiculid nymphs is seem-
ingly specialized, but again body shapes vary ^vithin the family
Trombidiidae and there is little to suggest what shape is primitive
for the trombiculids.

The specialized conditions for these characters are the alternative
ones given in the section on morphology. To repeat them here seems
unnecessary. Many of the specialized states are mentioned in the
following discussion.

It is more difficult to suggest primitive characters for the larval
stage. Few trombidiid larvae are known; most of the described
forms belong to the more highly developed trombidiids. The

Nymphs of the Trombiculidae 279

gnathosomal elements are particularly difficult, since both the chelic-
eral blade and the palpus seem specialized in larvae of both fam-
ilies. For the scutum a rectangular shape, an anteromedian projec-
tion, two anteromedian setae, unexpanded sensilla, and four mar-
ginal setae seem primitive. Possibly, undivided femora of the legs
is a primitive character but this conclusion is based entirely on com-
parisons within the family.

To return to the nymphal characters, it is possible to follow in a
general way some features of the evolutionary development of two
structural areas, the gnathosoma and the scutum. The characters
listed above as primitive are found among the genera of the sub-
family Leeuwenhoekiinae. Although there is some deviation in
hypostomal shape, shape of cheliceral elements, and presence of
eyes, the general facies for these areas in the Leeuwenhoekiinae is
primitive. The gnathosomal elements show some modifications in
most genera of Trombiculinae. Except for reduced palpal setation
and slight cheliceral modification in some species, the genera
Eiischongastia, Speleocola, and Tromhicula have a primitive condi-
tion of gnathosomal elements. Bkinkaartia differs only in the posses-
sion of additional accessory claws on the palpus in some species, a
situation common in the trombidiids. Greater modification is seen
in Neoschongosfia, where palpal claws are elongated, chelicerae
shortened, and hypostome blunt; Cheladonta has all gnathosomal
elements elongated. Finally, the genera Euschongastoides, Pseudo-
schongastia, and Walchia have distinctive chelicerae and markedly
modified hypostoma.

Scutal elements are also modified in the subfamily Trombiculinae.
These are discussed in detail in the section on moqohology. Most
noticeable specializations are the reduction in number of tectal setae,
shortening and thickening (in some) of the crista, development of
ridges and other structures in the sensillary area, modification or
reduction of the posterior apodeme, and modification of the sen-
silla. The genus Bkinkaartia, aside from the possession of a single
tectal seta and slight development of the sensillary area, has an
essentially primitive scutum. Development of the sensillary area
seems to proceed along two different lines among the remaining
genera. The sensillary areas of the genera Euschongastoides,
Fseudoschongastia, and Walchia are quite broadened. The remain-
ing genera have less broadened but seemingly more complicated
sensillary areas; those of Clieladonta and Neoschongastia each have
distinctive features not shared by Eiischongastia, Tromhicula, and
Speleocola.

280 The University Science Bulletin

From these considerations it is possible to divide the genera into
three levels of increasing specialization. The first level, the more
primitive state, includes genera of the subfamily Leeuwenhoekiinae.
The second level, an intermediate state, includes the genera Blanka-
artia, Euschongastia, Speleocola, and Trombicula. The third level,
the most specialized state, includes the genera CheJadonta, Neo-
sclwngastia, Euschongastoides, Pseudoschongastia, and Walchia. It
seems likely that these levels represent an evolutionary sequence.

Unfortunately it is not possible to consider more detailed rela-
tionships within the Leeuwenhoekiinae. Among the five genera
only eight species are known as nymphs and these appear to be but
slightly modified. Speculation as to their relationships would be
fruitless. Possibly, also, the habit of parasitism on vertebrates has
arisen more than once and a realistic evaluation must consider the
very similar forms in the Trombidiidae.

Blankaartia is the least modified of the intermediate genera and
might be considered halfway between the primitive forms and the
other intermediate ones. The other intermediate genera, Trom-
bicula, Euschongastia, and Speleocola appear closely related. The
genera Euschongastoides, Pseudoschongastia and Walchia are sim-
ilar and quite possibly represent a valid unit. While they are un-
hesitatingly considered specialized, it is difficult to avoid the im-
pression that in these genera the sensillary area of the scutum is
less modified than is that of the genera of the intermediate level.
It might be more reasonable to suggest that these three genera are
separately derived from the primitive forms, or at least that they are
an early branch from the intermediate group of genera. The other
specialized genera, Cheladonta and Neoschongastia seem distinctive.

Features of the larval stage essentially confirm this general se-
quence. The subfamily Leeuwenhoekiinae again clearly represents
the primitive state. A close relationship between larvae of genera
of the intermediate state appears. The specialized genera Chela-
donta, Neoschongastia, Euschongastoides, and perhaps Pseudo-
schongastia seem related to the intermediate genus Euschongastia
in features of scutal shape and nature of the sensilla. It has been
previously noted, however, that the character of the sensilla (ex-
panded vs. flagelliform) which separates the genera Euschongastia
and Trombicula has led to strange relationships.

While larva and postlarva are adapted to different sorts of exist-
ence, the divergence in form is not extreme when compared to
other animals with divergent phases of the life history (the holo-
metabolous insects, for instance). Although obviously modified,

Nymphs of the TROMBicuLroAE 281

the scutal and gnathosomal elements likely perform identical or
similar functions in the two stages. In these mites a possible
evolutionary mechanism is the transfer of a structural modification
from one stage to another stage. The sensillum of the larva of
Euschongastoides hoplai seems identical with the sensillum of the
nymph of that species; it is possible that in this case the larva has
acquired the sensillum of the nymph, since other similar larvae have
expanded sensilla. Sensilla may serve different purposes in the two
phases of the life history but it is possible that adaptive modifica-
tions in the nymph could be of advantage to the larva. No evidence
of other such transfers was noted. It is possible, however, that the
transfer of such preformed structures from one stage to another
differently adapted one could play an important role in the evolution
of these mites.

As more postlarvae are described these suggestions as to phy-
logeny will doubtless require modification. A productive study
would be the ecology and functional physiology of these mites, to
provide an indication of the specific uses of the structures here con-
sidered and thus a better understanding of the selective forces pro-
ducing them.

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1954. Notes on the taxonomy of trombiculid mites with description of a
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Borland, J. G.

1955. The genus Neotrambidium (Acarina: Trombidioidea ) in the United
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DOMROW, R.

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1947. A comer test for association. Ann. Math. Stat., vol. 18, pp.
495-513.

Philip, C. B., and Fuller, H. S.

1950. The harvest mites ("akidani") of Japan and the Far East and
their relationship to the autumnalis group of trombicuhd mites.
Parasit., vol. 40, pp. 50-57.

Nymphs of the Trombiculidae 283

Richards, W. S.

1950. The variation ot the British harvest mite. Parasit., vol. 40, pp.
105-117.

Sasa, M.

1953. Description of nymphs and adults of Japanese trombicuHd mites
reared in the laboratory. ( Studies on tsutsugamushi, part 54. )
Japanese Jour. Exp. Med., vol. 23, pp. 407-450.
Wharton, G. W.

1946. Observations on Ascoschongastia indica (Hirst, 1915) (Acarinida:
Trombiculidae). Ecol. Monographs, vol. 16, pp. 151-184.
Wharton, G. W., and Carver, R. K.

1946. Food of nymphs and adults of Neoschongastia indica (Hiist, 1915).
Science, vol. 104, pp. 76-77.
Wharton, G. W., and Fuller, H. S.

1952. A manual of the chiggers. Mem. Ent. Soc. Washington, no. 4,
pp. 1-185. Washington, D. C.
Wharton, G. W., and Hardcastle, A. B.

1946. The genus Neoscliongastia (Acarinida: Trombiculidae) in the
western pacific area. Jour. Parasit., vol. 32, pp. 286-322.
Wharton, G. W., Jenkins, D. W., Brennan, J. M., Fuller, II. S., Kohls,
G. M., and Philip, C. B.

1951. The terminology and classification of trombiculid mites (Acarina,
Trombiculidae). Jour. Parasit., vol. 37, pp. 13-31.

Willmann, C.

1941. Tromhicida minor Berl. 1905, typenart der gattung Trombicula
Berlese ( Trombidiidae, Acari). Zool. Anz., vol. 133, pp. 131-136.
Wolfenbarger, K. a.

1952. Systematic and biological studies on North American chiggers of
the genus Trombicula, subgenus Eutrombicula (Acarina, Trom-
biculidae). Ann. Ent. Soc. America, vol. 45, pp. 645-677.

Womersley, H.

1945. Acarina of Australia and New Guinea. The family Leeuwen-

hoekiidae. Trans. Roy. Soc. South Australia, vol. 69, pp. 96-113.
1952. The scrub-typhus and scrub-itch mites (Trombiculidae, Acarina)

of the Asiatic-Pacific region. Rec. South Australian Mus., vol. 10,

pp. 1-435.

284

The University Science Bulletin

Explanation of Figures 1-7
Tromhicula trisetica, nymph
Fig. 1. Body shape, dorsal view.
Fig. 2. Palpus, medial view.
Fig. 3. Cheliceral blade, lateral view.
Fig. 4. Scutum, dorsal view.
Fig. 5. Chelicera, lateral view.
Fig. 6. Cheliceral apodeme, lateral view.
Fig. 7. Posterior dorsal body seta.

a. palpus n.

b. chelicera , o.

c. scutum p.

d. trochanter q,

e. femur r.

f. genu s.

g. tibia t.
h. accessory claws u.
i. claw V.
j. tarsus w.
k. teeth of tectal margin x.
1. tectal seta
m. tectiun

sensillum
crista
carina
buUa

basal plate
sensiUary base
posterior apodeme
cheliceral blade
pseudochela
cheliceral base

area of articulation of chelic-
eral base with basis capituli

Nymphs of the Trombiculidae

285

Figures 1-7

286 The University Science Bulletin

Explanation of Figures 8-15

Features of Nymphs. (Not to same scale.)
Fig. 8. Euschdngastoides loomisi, hypostome.
Fig. 9. Trombicula montanensis, hypostome.
Fig. 10. Cheladonta ouachitensis, hypostome (lateral view).
Fig. 11. NeoschiJngastia brennani, tip of hypostome.
Fig. 12. Trombicula montanensis, genitalia.
Figs. 13-15. Trombicula splendens, variations in branching of genital setae.

Nymphs of the Trombiculidae

287

Figures 8-15

/iftN,

• ."•■>■

288 The University Science Bulletin

Explanation of Figures 16-29 j

Chelicerae of Nymphs. ( Not to same scale. )

Fig. 16. Chatia setosa, chelicera.

Fig. 17. Chatia setosa, cheliceral blade. j

Fig. 18. Leeuwenhoekia americana, chelicera, cheliceral blade, and

cheliceral apodeme. j

Fig. 19. Hannemania tnultifemorala, chelicera and cheliceral blade. ,

Fig. 20. H. eltoni, chelicera, cheliceral blade, and cheliceral apodeme. ;

Fig. 21. Trombicula alfreddugesi, chelicera and cheliceral blade. j

Fig. 22. Blankaartia alleei, chelicera and cheliceral blade. i

Fig. 23. Trombicula helkini, chelicera and cheliceral blade.

Fig. 24. T. splendens, chelicera and cheliceral blade. 1

Fig. 25. T. lipovsktjana, cheliceral blade. ;

Fig. 26. Euschongastia jonesi, chelicera and cheliceral blade. |

Fig. 27. E. setosa, chelicera and cheliceral apodeme.

Fig. 28. E. pipistrelli, cheliceral blade. i

Fig. 29. E. setosa, cheliceral blade. \

Nymphs of the Trombiculidae

289

Figures 16-29

10-8920

290 The University Science Bulletin

Explanation of Figures 30-44
Chelicerae of Nymphs. (To same scale except where indicated.)
Euschdngastoides loomisi, chelicera ( not to scale ) .
E. hoplai, chelicera (not to scale).

Fseudoschongastia htingerfordi, chelicera and cheliceral blade.
P. fameri, chelicera and cheliceral blade.
Acomatacarus plumosus, chelicera and cheliceral blade.
Walchia americana, chelicera and cheliceral blade.
Whartonia senase, chelicera and cheliceral blade.
Acomatacarus arizonen^is, chehcera and cheliceral blade.
Trombicula merrihewi, chehcera and cheHceral blade.
Speleocola tadaridae, chelicera and cheliceral blade.
Neoschongastia americana, chelicera and chehceral blade.
N. brennani, chelicera and cheliceral blade.
Cheladonta micheneri, chelicera.
C. micheneri, chehceral blade.
C ouachitensis, chehceral blade.

Fig.

30.

Fig.

31.

Fig.

32.

Fig.

33.

Fig.

34.

Fig.

35.

Fig.

36.

Fig.

37.

Fig.

38.

Fig.

39.

Fig.

40.

Fig.

41.

Fig.

42.

Fig.

43.

Fig.

44.

Nymphs of the Trombiculidae

291

Figures 30-44

292

The University Science Bulletin

Explanation of Figures 45-59

Chelicerae of Nymphs. ( To same scale. )

Fig. 45. Euschdngastia trigenuala, chelicera and cheliceral bladt'.

Fig. 46. E. criceticola, chelicera and cheliceral blade.

Fig. 47. E. pipistrelli, chelicera and cheliceral blade.

Fig. 48. E. peromysci, chelicera and chehceral blade.

Fig. 49. E. diversa, chelicera and cheliceral blade.

Fig. 50. E. cynomyicola, chelicera and cheliceral blade.

Fig. 51. Trombicula tvhartoni, chelicera, cheliceral blade, and cheliceral
apodeme.

Fig. 52. T. autumnalis, chelicera and cheliceral blade.

Fig. 53. T. sylvilagi, chelicera and cheliceral blade.

Fig. 54. T. montanensis, chelicera and cheliceral blade.

Fig. 55. T. myotis, chelicera and cheliceral blade.

Fig. 56. T. kansasensis, chelicera and chehceral blade.

Fig. 57. T. gurneyi, chelicera, cheliceral blade, and cheliceral apodeme.

Fig. 58. T. omata, chelicera and cheliceral blade.

Fig. 59. T. kardosi, chelicera, cheliceral blade and cheliceral apodeme.

Nymphs of the Trombiculbdae

293

FiGUBEs 45-59

294 The University Science Bulletin

Explanation of Figures 60-73 i

Medial Views of Palpi of Nymphs.
(To same scale except where indicated.)

Fig. 60. Whartonia senase. i

Fig. 61. Hannemania eltoni (not to scale). [

Fig. 62. Leeuwenhoekia americana.

Fig. 63. Blankaartia alleei (not to scale). |

Fig. 64. EuscJwngastia jonesi. I

Fig. 65. E. setosa. '

Fig. 66. E. peromijsci.

Fig. 67. E. cynomijicola.

Fig. 68. E. trigenuala.

Fig. 69. Trombicula giimeyi.

Fig. 70. Euschongastia criceticola.

Fig. 71. Neoschongastia americana.

Fig. 72. N. hrennani.

Fig. 73. Tr&mbicula montanensis.

Nymphs of the Trombiculidae

295

Figures 60-73

296 The University Science Bulletin

Explanation of Figures 74-83
Medial Views of Palpi of Nymphs. ( To same scale. )
Fig. 74. Trombiculu alfreddugisi.
Fig. 75. T. splendens.
Fig. 76. T. lipovskyana.
Fig. 77. T. kansasensis.
Fig. 78. T. kafdosi.
Fig. 79. T. belkini.

Fig. 80. T. whartoni (includes apical nude setae on tarsus).
Fig. 81. r. my Otis.
Fig. 82. T. autumnalis.
Fig. 83. T. lipovskyi.

Nymphs of the Trombiculidae

297

Figures 74-83

298 The University Science Bulletin

Explanation of Figures 84-91

Medial Views of Palpi of Nymphs. (To same scale.)

Walchia americana.

Cheladonta micheneri.

Speleocola tadaridae.

Euschdngastoides hoplai.

Acamatacanis plumosus (including dorsal genual, dorsal femoral,
and lateral femoral seta).

Fig. 89. Trombicula merrihewi.
Fig. 90. Pseudoschongastia fameri.
Fig. 91. P. hungerfordi.

Fig.

84.

Fig.

85.

Fig.

86.

Fig.

87.

Fig.

88.

Nymphs of the Trombtculidae

299

Figures 84-91

300 The University Science Bulletin

Explanation of Figures 92-101

Palpal Tibiotarsi of Nymphs. (To same scale.)

Fig. 92. Chatia setosa, lateral view.

Fig. 93. C. setosa, medial view.

Fig. 94. Whartonia senase, lateral view.

Fig. 95. W. senase, medial view.

Fig. 96. Leeuwenhoekia americana, lateral view.

Fig. 97. L. americana, medial view.

Fig. 98. Hannemania eltoni, lateral view.

Fig. 99. H. eltoni, medial view.

Fig. 100. Neoschongastia brennani, lateral view.

Fig. 101. N. brennani, medial view.

Nymphs of the Trombiculidae

301

Figures 92-101

302 The University Science Bulletin

Explanation of Figures 102-117

Palpal Tibiotarsi of Nymphs. (To same scale.)

Fig. 102. Euschongastia setosa, lateral view.

Fig. 103. E. setosa, medial view.

Fig. 104. E. peromysci, lateral view.

Fig. 105. £. peromysci, medial view.

Fig. 106. Trombicula myotis, lateral view.

Fig. 107. T. myotis, medial view.

Fig. 108. E. jonesi, lateral view.

Fig. 109. E. jonesi, medial view.

Fig. 110. Trombicula alfreddugesi, lateral view.

Fig. 111. T. alfreddugesi, medial view.

Fig. 112. T. gumeyi, lateral view.

Fig. 113. T. gumeyi, medial view.

Fig. 114. T. kansasensis, lateral view.

Fig. 115. T. kansasensis, medial view.

Fig. 116. T. lipovskyi, lateral view.

Fig. 117. T. lipovskyi, medial view.

Nymphs of the Trombiculidae

303

Figures 102-117

304 The University Science Bulletin

Explanation of Figures 118-137

Palpal Tibiotarsi of Nymphs. (To same scale.)

Fig. 118. Walchia americana, lateral view.

Fig. 119. W. americana, medial view.

Fig. 120. Euschongastia trigenuala, lateral view.

Fig. 121. E. trigenuala, medial view.

Fig. 122. Trombictila montanensis, lateral view.

Fig. 123. T. montanensis, medial view.

Fig. 124. Pseudoschongastia farneri, lateral view.

Fig. 125. P. farneri, medial view.

Fig. 126. Euschongastoides hoplai, lateral view.

Fig. 127. E. hoplai, medial view. 1

Fig. 128. SpeleocoJa tadaridae, lateral view.

Fig. 129. S. tadaridae, medial view. j

Fig. 130. Euschongastia criceticola, lateral view. i

Fig. 131. E. criceticola, medial view.

Fig. 132. Tromhicula crossleyi, lateral view. j

Fig. 133. T. crossleyi, medial view.

Fig. 134. Acomatacanis plumosus, lateral view.

Fig. 135. A. plumosus, medial view.

Fig. 136. Tromhicula merrihewi, lateral view.

Fig. 137. T. merrihewi, medial view.

Nymphs of the Trombiculidae

305

Figures 118-137

306 The University Science Bulletin

Explanation of Figures 138-147
Scuta of Nymphs. (To same scale except where indicated.)
Fig. 138. Acamatacarus plumosus.
Fig. 139. Hannemania cltoni.
Fig. 140. Leeuwenhoekia amcricana.
Fig. 141. Cluitia setosa.

Fig. 142. Acomatacarus arizonensis (not to same scale).
Fig. 143. Whartonia senase.

Fig. 144. Pseudoschongastia Imngerfordi (sensillum).
Fig. 145. P. farneri (not to same scale).
Fig. 146. Euschongastoides hopJai.
Fig. 147. Walchia americana (not to same scale).

Nymphs of the Trombiculidae

307

FiGUBEs 138-147

308 The University Science Bulletin

Explanation of Figures 148-156

Scuta of Nymphs. (To same scale except where indicated.)
Fig. 148. Blankaartia velascoi (not to same scale).
Fig. 149. Neoschongastia brennani.
Fig. 150. Euschongastia criceticola.
Fig. 151. E. trigenuala.
Fig. 152. E. peromysci.
Fig. 153. E. jonesi.
Fig. 154. Cheladonta micheneri.
Fig. 155. C. ouachitensis (sensillum).
Fig. 156. Euschongastia setosa.

Nymphs of the Trombiculidae

309

FiGiTRES 148-156

310 The University Science Bulletin

Explanation of Figures 157-168

Scuta of Nymphs, (to same scale except where indicated

Fig. 157.

Trombicula autumnaUs (not to same scale).

Fig. 158.

T. lipovskyi.

Fig. 159.

T. whartoni.

Fig. 160.

T. sylvilagi (sensillum).

Fig. 161.

T. kardosL

Fig. 162.

T. ornata.

Fig. 163.

T. crossleyi (sensillum).

Fig. 164.

T. gurneyi.

Fig. 165.

T. merrihewi.

Fig. 166.

T. myotis.

Fig. 167.

T. mantanensis.

Fig. 168.

T. alfreddugesi.

Nymphs of the Trombiculidae

311

Figures 157-168

312 The University Science Bulletin

Explanation of Figures 169-183 \

Features of Nymphs. (Not to same scale.) i

Fig. 169. Blankaartia velascoi, lateral view of palpal tibiotarsus.

Fig. 170. B. velascoi, medial view of palpal tibiotarsus.

Fig. 171. B. alleei, lateral view of palpal tibiotarsus.

Fig. 172. B. alleei, medial view of palpal tibiotarsus.

Fig. 173. Speleocola tadaridae, scutum. '

Fig. 174. Trombicula kardosi, lateral view of palpal tibiotarsus.

Fig. 175. T. kardosi, medial view of palpal tibiotarsus.

Fig. 176. T. splendens, sternum. <

Fig. 177. Neoschdngastia americana, sternum. |

Fig. 178. Hannemania multifemorala, sternum. •

Fig. 179. Euschongastoides haplai, sternum. I

Fig. 180. Neoschdngastia americana, scutum. i

Fig. 181. Trombicula merrihewi, sternum. \

Fig. 182. Whartonia senase, sternum. I

Fig. 183. Cheladonta micheneri, sternum. I

Nymphs of the TROMBicuLroAE

313

Figures 169-183

314 The University Science Bulletin

Explanation of Figures 184-199
Features of Nymphs. (Not to same scale.)
Fig. 184. Cheladonta micheneri, tibia and tarsus of leg I.
Fig. 185. Leeuwenhoekia americana, tibia and tarsus of leg I.
Fig. 186. Euschongastoides hoplai, tibia and tarsus of leg I.
Fig. 187. Whartonia senase, tibia and tarsus of leg I.
Fig. 188. Walchia americana, tibia and tarsus of leg I.
Fig. 189. Acomatacarus plumosus, tibia and tarsus of leg I.
Fig. 190. Chatia setosa, tibia and tarsus of leg I.
Fig. 191. Trombicula alfreddugesi, tibia and tarsus of leg I.
Fig. 192. Euschongastia trigenuala, tibia and tarsus of leg I.
Fig. 193. Neoschongastia americana, lateral view of palpal tibiotarsus.
Fig. 194. N. americana, medial view of palpal tibiotarsus.
Fig. 195. Cheladonta micheneri, lateral view of palpal tibiotarsus.
Fig. 196. C micheneri, medial view of palpal tibiotarsus.
Fig. 197. Leeuwenhoekia americana, anus.

Fig. 198. Trombicida tricetica, dorsal view of basis capituli and h\postome
(palpi still articulated at sides but chelicerae removed).
Fig. 199. Trombicula splendens, anus.

Nymphs of the TROMBicuLroAE

315

Figures 184-199

316 The University Science Bulletin

Fig.

200.

Fig.

201.

Fig.

202.

Fig.

203.

Fig.

204.

Fig.

205.

Fig.

206.

Fig.

207.

Fig.

208.

Fig.

209.

Fig.

210.

Explanation of Figures 200-210

Features of Nymphs. ( Not to same scale. )

Cheladonta ouachitensis, dorsal outline of body.
Neoschongastia americana, dorsal outline of body.
Trombicula omata, dorsal outline of body,
Walchia americana, dorsal outline of body.
Hannemania dunni, posterior body seta.
Acomatacarus plumosus, dorsal outline of body.
Blankaartia velascoi, posterior body seta.
Acomatacarus plumosus, posterior body seta.
Blankaartia alleei, posterior body seta.
Acomatacarus arizonensis, posterior body seta.
Chatia setosa, posterior body seta.

Nymphs of the Trombiculidae

317

Figures 200-210

209

318 The University Science Bulletin

Explanation of Figures 211-225 i

Posterior Body Setae of Nymphs. (To same scale except where indicated.) j

Fig. 211. Euschbngastia peromysci (tip) (Tennessee variant). i

Fig. 212. E. cynomtjicola (tip).

Fig. 213. £. setosa (tip). j

Fig. 214. E. peromysci (tip) (Kansas variant). '

Fig. 215. E. diversa (tip). ;

Fig. 216. £. jonesi (tip). i

Fig. 217. E. criceticola.

Fig. 218. Pseudoschongastia hungerfordi.

Fig. 219. Euschongastoides hoplai. ..

Fig. 220. Euschongastia trigenuala. i

Fig. 221. Cheladonta ouachitensis. '^

Fig. 222. Neoschongastia americana (long seta). 1

Fig. 223. Euschongastoides loomisi ( tip ) . 4

Fig. 224. Neoschongastia americana (short seta). t!

Fig. 225. Euschongastia pipistrelli (not to same scale). 1

Nymphs of the Trombiculidae

319

Figures 211-225

220

225 \

320 The University Science Bulletin

Explanation of Figures 226-240 ,

Posterior Body Setae of Nymphs. (To same scale except where indicated.) i
Figs. 226-227. Tromhicula alfreddugesi.

Fig. 228. r. splendens (tip). j

Fig. 229. r. helkini. \
Fig. 230. T. fitchi (not to scale).

Fig. 231. T. myotis. :
Fig. 232. T. merrihewi.

Fig. 233. Speleocola tadaridae. i

Fig. 234. Whartonia senase. \

Fig. 235. Walchia americana. ;
Fig. 236. Tromhicula autumnalis.

Fig. 237. T. sylvilagi. f

Fig. 238. T. gurneyi. '

Fig. 239. T. montanensis. ,

Fig. 240. T. lipovskyi. I

Nymphs of the Trombiculidae

321

Figures 226-240

226

11—8920

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 7

Concerning the Genus Ventidius and Five New Species

( Heteroptera, Gerridae ) '^

Herbert B. Hungerford and Ryuichi Matsuda

Abstract — This paper contains the descriptions of five new species: Ven-
tidius malayensis and \^. chinai from Malay peninsula, V. usingeri and V^.
werneri from the Philippines, and V. (Ventidioides) kttiterti subg. et sp. nov.
from Burma. It also presents a key to all the species of the genus.

The genus Ventidius was described by Distant for a small Halobatine gerrid
from Travancore, India, which he named Ventidius aquarius (Ann. Mag. .\at.
Hist, ser. 8, vol. 5, 1910, p. 1.50). In the Fauna of British India, Rhynchota
V, Heteroptera, appendix p. 157 (1910) Distant figured both winged and
wingless individuals of the same species. In 1918 Pai\a described V. distunti
from Yawnghwe State (now in East Burma) ( Rec. Ind. Mus., vol. 14, p. 25, pi.
8, fig. 4). In 1930 Esaki described V. henrtji from Ceylon (Jour. Fed. Malay
Mus., vol. 16, p. 18, pi. I and 2). In 193'3 Lundblad described V. modulatus
from Java (Arch. Hydrobiol. Suppl. Bd. 12, Tropische Binnengewiisser 4,
s. 399). While all of these authors gave pictures of their species only Lund-
blad pictured the male genitalia of his species.

We thought it would be a simple matter to determine the specimens we
have from Burma and Philippines. Howcxer, we found that the males of
our Burmese species possessed two distinctive characters not mentioned in
previous descriptions and this required the re-e.\aniination of the types and
the assistance of the entomologists in charge of them. We are therefore
grateful to Dr. W. E. China and Mr. R.- J. Izzard of the British Museum for
re-examining the types of V. aquarius and V^. hetiryi, to Dr. A. P. Kapur
of the Zoological Survey of India, Calcutta, for answering our inquiries about
T. distanti Pai\a, and to Dr. O. Lundblad of Stockholm, Sweden, for giving
us additional information about his V. modulatus. With their help we have
attempted to place the four described species, to describe fi\e new species,
and to propose a new subgenus to include two species that ha\e characters
not previously noted in the genus.

In course of our study we ha\c found that the sutures occurring in the
thorax and basal abdominal tergites are important in separating the species.

* Contribution No. 1011 from the Department of Entomology, Universit>' of Kansas.
This paper is a by-product of a project by a grant from the National Science Foundation.

(323)

324 The University Science Bulletin

To facilitate the understanding of the description of these sutures in the text
we have diagrammatically shown a hypothetically primitive Ventidius species
possessing all sutures with which we are concerned on plate 3, figs. 1, 2.

Ventidius aquarius Distant
(PI. 1, figs. 1, 2)

1910 Distant, W. L. Ann. Mag. Nat. Hist., ser. 8, vol. 5, p. 150 (described
Ventidius with aquarius as the type species from Travancore, S. India).

1910 Distant, W. L. Fauna Brit. Ind. Rhynchota 5, Heteroptera: appendi.x
pp. 157-158, fig. 84 (gives description and figures of winged and wingless
individuals ) .

1911 Bergroth, E. Ann. Soc. Ent. Bclgique, tome 60, p. 186 (considered
Ventidius as a synonym of Metrocoris Mavr).

1928 Esaki, T. Ann. Mag. Nat. Hist., ser. 10,' vol. 2, p. 572 (recorded from
South India).

1928 Dover, C. Treubia, Dcel 10, AH. 1. Biz. 69 (says V. distanti PaiNa is a
synonym of V. aquarius).

1930 Esaki, T. Jour. Federated Malay Mus., vol. 16, p. 18, pis. 1 and 2 (says
"a cotype of this species, a single apterous male* from Pallode near
Trivandrum, Travancore, is now to be found in the British Museum. Its
length of body is 3.5 mm., and ratio of its antennal segments:: 18:9:9:7.
Then he gives a new record "1 9 Sunpai Ampang, Selangor, Federated
Malay State, Aug. 15, 1926 C. Dover. ' We found this to bo another
species.

1933 Lundblad, O. Arch. Hydrobiol. Suppl. Bd. 12, Tropische Binnenge-
wasser, 4, S. 399.

Since we have not seen the type we quote Distant's description
below and have copied the pictures given by him on pi. 1, figs. 1, 2.

"Macropterous form: Head and pronotum ochraceous, the latter
with a large cruciform fascia and the sublateral and apical margins
black; eyes black; hemelytra black; body beneath ochraceous; legs
and apex of rostrum black, bases of anterior femora ochraceous;
acetabula and coxae spotted with black; structural characters as
in generic diagnosis.

"Apterous form. Body above ochraceous; pronotum with a nar-
row sublateral fascia, angulated and reaching margin at humeral
angle, and two spots at centre of posterior margin continued across
mesonotum, black, other markings generally as in macropterous
form; abdomen above ochraceous, with the segmental margin black.

"Length, macropt. form 5 mm., apt. form 4*2 mm.

"Hah. Travancore, Pallode, 20 miles N. E. of Trivandrum (An-
nandale) 'On roadside jungle stream (N. Annandale)'." An apter-
ous female specimen from Travancore is in the British Museum.

This species is known only from the types from Travancore, South
India. Dover considered V. distanti to be a synonym of V. aquarius,

* In 1958 Dr. China wrote us that the single cotype of V. aquarius in the British
Museum is a female, apterous, and lacks entirely the intersegm'. ntal suture between the meso-
and metanota.

Concerning the Genus Ventidius

325

but we believe he was in error. Dr. A. P. Kapnr wrote to us that
they have a single female apterous specimen of this species from
Travancore. A more satisfactory description of this species awaits
the capture of more specimens from South India. The specimens
reported by Esaki from the Malay Peninsula (1930) are of a new
species described below.

Ventidius malaijensis sp. nov.
(PI. 1 fig. 8, PI. 3 figs. 3, 5, 12)
Size: Apterous male 3.99 mm. long; 2.94 mm. wide across meso-
acetabula. Apterous females 3.57-4.1 mm. long; 2.94-3.05 mm. wide
across mesoacetabula.

Color: Shining testaceous, with dark brown to black markings
as shown in the figures on plate 2. Antennae except base of first
segment and legs, except basal half of front femur, black. Venter
pale.

Striicttiral characteristics: Relative lengths of antennal segments
in male: lst:2nd:3rd:4th: :90:60:32:40; in female: 72:35:27:27.
Male with a tuft of hairs near distal end of third segment. Anterior
margin of head somewhat produced beyond eyes which are sloping
and overlap lateral margins of prothorax and anterolateral angles
of mesothorax. Prothorax short, about half as long as head, with
anterior and posterior margins medially concave. Mesonotmn
swollen. Intersegmental suture between meso- and metanota faint
to obscure. Metanotum somewhat declivent; lateral suture of
metanotum not reaching the intersegmental suture anteriorly; lower
half of hind margin of metacetabula nearly transverse and l)road;
omphalium present. Anterior margin of first tergite lost. Con-
nexivum broad. Last abdominal ventrite as long as preceding
segments taken together in both sexes. Front tibia somewhat
swollen in the middle, claws arise from near base of second seg-
ment instead of near its middle as in V. aquarius and V. usingeri.

Relative lengths of leg segments of a male.

Femur

Tibia

First
tarsal

Second
tarsal

Total
of tarsal
segments

Front leg

225
640
690

205
390
360

Middle leg

Hind leg

?
110

326 The University Science Bulletin

Paramere small and symmetrical as in V. usingeri n. sp. but a little
different in shape. (See pi. 3 fig. 12).

Location of types: The male holotype and the female allotype
bear the label "Malay Peninsula, Selangor, F. M. S. Sungai Ampang,
Aug. 15, 1926 C. Dover, Ex. Coll. F. M. S. Museum." The male
holotype is also labelled '^Vcntidius ciqttarius Dist. det. by Teiso
Esaki," and the female allotype bears also the label "Metrocoris
(Uf minus Dist. det. Dover 26." A female paratype is labelled "Malay
Penin. Upper Perak, Lenggong Surface of irrigation channels fast
water I. H. N. E. 22 X 5— XI 1926 C. Dover" and on the reverse side
"Ex. coll. F. M. S. Museum 5 ." All the types are preserved at the
British Museum.

Ventidius usingeri sp. nov.
(PI. 2figs. 7, 8, PI. Sfigs. 4, 10)

Size: Winged male 4.2 mm. long; 2.06 mm. across humeri; 2.48
mm. across mesoacetabula. Winged female 4.62 mm. long; 2.1 mm.
across humeri; 2.56 mm. across mesoacetabula. Wingless female
3.68 mm. long; 2.73 mm. across mesoacetabula.

Color: Winged form pale yellowish with dark hemelytra, pronotal
black marking as shown on pi. 2. Apterous female with black
markings as shown on pi. 2, and the pale caudal lobe on metanotum
broadly rounded behind instead of having an angular projection
as in V. aquarius. The black markings, under certain reflections,
appear as iridescent green.

Structural characteristics: Relative lengths of antennal segments
in male: 1st: 2nd: 3rd: 4th: :75:53:38:28; in female: : 75: 43: 35: 26.
In winged forms eyes overlapping lateral region of prothorax, not
quite reaching propleural caudal margin. Caudal tip of pronotum
surpassing a line drawn between metathoracic spiracles; pale
median longitudinal line of pronotum bordered with dark lines
gives an appearance of being a carina. In wingless forms eyes over-
lap lateral margins of prothorax and anterolateral angles of meso-
thorax. Anterior and posterior margins of pronotum medially con-
cave. Intersegmental suture between meso- and metanota distinct
in one apterous female. Lateral suture of metanotum not reaching
intersegmental suture between meso- and metanota. Anterior
margin of first abdominal tergite obscure. Lower half of hind
margin of metaacetabula nearly transverse and broad. Last ab-
dominal ventrite in both sexes as long as all preceding segments
together. Parameres small and nearly symmetrical, only half as
long as those in V. distanti which is a much smaller species. Front

Concerning the Genus Ventidius

327

femur of male not swollen, very slightly curved and without a
tubercle on ventral side. First tarsal segment of middle leg curved
and ventrally hairy at base in both sexes. The male lacks meso-
sternal tubercle.

Relative lengths of leg segments of a i

male.

Femur

Tibia

First
tarsal

Second
tarsal

Total
of tarsal
segments

Front leg

210
530
575

178
330
240

10
167

53
26
37

Middle leg

193

Hind leg

Location of types: The holotype is a winged male bearing the
labels "Los Banos P. I. VII-17-19.36;' and "R. L. Usinger collector,"
and belongs to the Usinger collection. The allotype is a winged fe-
male bearing the labels "C N H M Philippines Zool. Exped. ( 1946-
1947)" and "Borungkot, Upi, Cotabato Province, Mindanao 1500
ft. '47." This belongs to the Chicago Natural History Museum.
A wingless paratype female specimen came from the same place
as the allotype and is in the Francis Huntington Snow museum.
University of Kansas.

Comparatwe notes: This species appears very much like V.
(Kitiarius, which was described from Travancore, South India.
The shape and size of parameres are as in V. nuilayensis sp. nov.,
but the relative lengths of antennal segments are different and the
male of V. nialayensis has a tuft of hairs near the distal end of the
third antennal segment, which is lacking in this species. As in
V. ciquarius the front leg has the claws arising from near the middle
of the second tarsal segment.

Ventidius henry i Esaki
(PI. 1 figs. 3, 4, PI. 2 fig. 6, PI. 3 fig. 8)

1928 Ventidius henrui Esaki, T. Ann. Mag. Nat. Hist. ser. 10, \-()l. 2. pp. 509-
511 ( described from Kitulgala, Ceylon. 1^,39 9 ).

1933 Ventidius henriji Lundblad, O. Arch. Hydrobiol. Suppl. 12, Tropischc
Binnengewiisser 4, S. 372 (records Ceylon).

Dr. W. E. China of the British Museum kindly answered our
inquiry by reporting that they have one male (type) and one
female from Ceylon, and that the mesosternum in the male speci-
men is spoiled by the pin, and also that the anterior femora lack
the small tubercle present in V. kuiierti sp. nov.

328 The University Science Bulletin

We are grateful also to Dr. P. E. P. Deraniyagala, Director of
the National Museum, Colombo, Ceylon, for sending us in alcohol
15 adults and 11 nymphs taken from Kitulgayala in Sabaragamuwa
Prov., Ceylon, in December 1934. These are therefore possibly
from the type locality but not of the type series and enable us to
add to the kownledge of this species, for Esaki knew only apterous
specimens and did not describe the male parameres.

Size: Winged female: 2.73 mm. long; across humeri 1.5 mm.
wide; across mesoacetabula 1.8 mm. wide. Wingless female: 2.5-
2.94 mm. long; across mesoacetabula 1.78-1.89 mm. wide. Wingless
males: 2.3-2.4 mm. long; across mesoacetabula 1.43-1.5 mm. wide.
Although Esaki gives the male as 2.4 mm. long and 1.8 mm. wide,
and the female as 2.2 mm. long and 1.6 mm. wide in our speci-
mens the females are all larger than the males.

Color: Wingless forms have the pattern as shown on pi. 1 fig. 4.
Surface not shining, finely pilose. Pale band on head usually red-
dish, the other spots stramineous. Pronotum black. Sides of
mesothorax with longitudinal dark brown to black band behind
eyes. Venter pale. Antennae except base, legs and beak brown.
Winged specimen which is teneral dark brown instead of black in
ground color on upper surface; head wtih the characteristic reddish
band and pronotum with stramineous spots.

Structural characteristics: Relative length of antennal segments in
male: 1st: 2nd: 3rd: 4th: :46:20:18:16; in female: 45:18:20:18 (with
some variations). While Esaki gives the formula for the species as
50:20:20 he says "second, third and fourth segments nearly equal in
length." Eyes in apterous forms overlapping lateral margins of pro-
thorax and anterolateral angles of mesothorax. Anterior margin of
pronotum between eyes nealy straight, posterior margin concave.
Intersegmental suture between meso- and metanota faint to obscure.
Lateral suture of metanotum not reaching intersegmental suture
anteriorly. Anterior margin of first tergite faint to obscure. Lower
half of caudal margin of metaacetabula oblique, its lower angle
nearly pointed. Omphalium visible. Last abdominal ventrite short,
scarcely as long as total length of all abdominal ventrites that pre-
cede it and much shorter than the venter of eighth segment in male.
Eighth ventrite a little longer than all segments together that precede
it in male. Last ventrite in female short, not as long as all preceding
segments together. Front femur without protuberance in male and
mesosternum lacks a protuberance in male. Parameres as shown on
pi. 3 fig. 8. They are symmetrical.

Concerning the Genus Ventidius

329

Relative lengths of leg segments of a male.

Femur

Tibia

First
tarsal

Second
tarsal

Total
length

of
tarsus

Front leg . .

135
405
450

120
219
145

10.2

116.7

28.0

37.8
24.3
29.0

Middle leg

Hind leg

141
57

Location of types: The types are in the British Museum. The
specimens studied in this work are in the Francis Huntington Snow
Museum, University of Kansas.

Ventidius modulatus Lnndblad

(PI. 1, fig. 7)

1933 Ventidius modulatus Lnndblad, O. Arch. Hvdrobiol. Suppl. Bd. 12. Trop-
ische Binnengewiisser 4, S. 399-401, Taf. 12, and fig. 128.

Dr. Lnndblad gave an excellent description and good illustration
of this species from West Java. We give here some statements from
his description.

Size: About 2.5 mm. long.

Color: Ground color lemon yellow. Head with a small brown
fleck in front. Behind each eye on pro-, meso-, and metanotum with
a fairly broad brown longitudinal band. Behind the boundary of
meso- and metanota with a lateral band of brown color, which may
be free at its end or be joined to a dark fleck on metaacetabula. Base
of antennae and of front leg yellowish. Anterior abdominal tergites
dark as shown on plate, or have yellowish triangular flecks in the
middle. Posterior tergites and the entire venter alway.s yellow.
Connexivum largely yellow.

Stnictura] characteristics: Relative lengths of antennal segments:
1st: 2nd: 3rd: 4th::53:27.6:20.3:19. The suture between meso- and
metanota distinct, also the one between metanotum and first abdom-
inal tergite distinct. First three abdominal tergites confluent with-
out distinct sutures (with distinct sutures in Taf. 12). Male para-
mere relatively large.

Location of types: Described from four males taken from West
Java, Stausee Tjigombong, south of Buitenzorg, 500 in. above sea
level. 17. IX. 1928. Types are in the National Museum. Stock-
holm, Sweden.

330

The University Science Bulletin

Ventiditis werneri sp. nov.
(PI. 2figs. 4, 5, PI. 3, fig. 11)

Size: Wingless male 2.2 mm. long; 1.55 mm. wide. Wingless
female 2.3 mm. long; 1.68 mm. wide.

Color: Black with whitish or yellowish white figures as shown
on plate 2. Head mostly whitish, with a black spot on vertex and
a black band along inner margin of each eye. Antennae black
except base of first segment. Pronotum black laterally, with a
transverse triangular whitish spot, its apex almost reaching anterior
margin of pronotum. The remaining thoracic dorsum with a large
conspicuous whitish spot that is broad at its apex and broadly bi-
lobed caudally; the large whitish spot on metanotum broadly con-
tinuous with mesonotal whitish spot. Mesothoracic pleuron behind
eye somewhat embrowned but lacking usual brown or black band.
Metaacetabular elongate whitish spot with its lower margin with a
dentate projection before middle and a lobate projection at caudal
angle. Abdominal tergites black, with last five or six tergites of fe-
male with median pale spots or bands that may cover most of the
tergites; those of male mostly black. Connexival segments of female
whitish; last five segments of male whitish. Legs black except base
of front femora.

Structural characteristics: Relative lengths of antennal segments
in male: 1st: 2nd: 3rd: 4th:: 48:22:19:20, those of female :: 43: 19: 21:
19. Eyes in wingless forms overlapping lateral margins of prothorax
and anterolateral angle of mesothorax. Anterior and posterior
margins of pronotum concave. Mesonotum swollen. Metanotum
more or less declivent. Intersegmental suture between meso- and
metanota faint to obscure. Lateral suture of metanotum not quite
reaching intersegmental suture between meso- and metanota.
Caudal margin of metaacetabula oblique and lower angle nearly
pointed. Front femora slightly curved in basal half; that of male
not thicker than that of female and without the tubercle charac-

Relative lengths of leg segments of a male.

Femur

Tibia

First
tarsal

Second
tarsal

Total
of tarsal
segments

Front leg

140
375
360

114
190
150

115

38
20
30

Middle leg

135

Hind leg

Concerning the Genus Ventidius

331

teristic of V. kuiterti. Male also lacks tubercle on mesosternum.
First tarsal segment of middle leg curved, thickened and ventrally
hairy at base in both sexes.

Location of types: Described from one male and two females
labelled "CNHM Philippines Zool. Exped. (1946-47). F. G. Wer-
ner, Puerto Princesa, Palawan Is. Sea level, March (1947)." Holo-
type and allotype are in the Chicago Natural History Museum,
and a female paratype is in the Francis Huntington Snow Museum,
University of Kansas.

Comparative notes: This species, which is about the same size
as V. distanti, has larger pale spots on pronotum and the mesonotal
spot is much broader anteriorly. The male lacks the tubercle of
front femur and the mesosternal tubercle.

Ventidius chinai sp. nov.
(PI. 1 figs. 5, 6, PI. 3 fig. 9)
Size: Wingless males 2.56-2.73 mm. long; across mesoacetabula
1.89 mm. wide. Wingless female 2.94 mm. long; 2.18-2.27 mm.
wide across mesoacetabula.

Color: Body pale stramineous. Antennae and legs, except base
of first antennal segment and of front femur, black. If any dark
spots occur on the body they are on sides of notum and base of
abdomen.

Structural characteristics: Relative lengths of antennal segments
in male: 1st: 2nd: 3rd: 4th : :56:28:20:20; those in a female ::48:22:
26:?. Eyes in wingless forms overlapping lateral margins of pro-
thorax and anterolateral angles of mesothorax. Anterior and pos-
terior margins of pronotum medially concave. Mesonotimi some-
what swollen. Intersegmental suture between nieso- and metanota
obscure. Metanotinu slightly declivent; lateral suture of metanotum
not reaching intersegmental suture between meso- and metanota.
Anterior margin of first tergite distinct; basal abdominal tergites
produced anteriorly, anterior margins obliterated medially on sec-
Relative lengths of leg segments of a male.

Femur

Tibia

First
t arsal

Second
tarsal

Total

of
tarsus

Front leg

Middle leg

Hind leg

150
450
450

135
246
200

7.8

1 29 (1

25.0

37 2
20 4
30.0

150

332 The University Science Bulletin

ond and third; last abdominal ventrite as long as all the preceding
segments together in both sexes. Connexivum rather broad. Hind
margin of metaacetabula oblique and lower angle nearly pointed.
Front femur of male without protuberance and not thicker than
that of female. Mesosternum in male lacks protuberance.

Male parameres are symmetrical and of characteristic shape; distal
end broadened and appears lobed in certain views. (Pi. 3, fig. 9.)

Location of types: Described from two wingless males and two
wingless females bearing the label "Malay Peninsula, Selangor,
F. M. S. Kajang Sungei Lang Feb. 12, 1927," on the reverse side "C.
Dover, Coll. F. M. S. Museum." Thev also bear another label "Ven-
tidiiis sp. no v. det. W. E. China 1934." The holotype male, allotype
female and two paratypes are in the British Museum.

Comparative notes: The surface of this pale species is shining and
nearly devoid of hairs. It is smaller than V. aquarius.

Ventidiiis distanti Paiva
(PI. 2, fig. 1)

1918 Ycntidius disanti Paiva, C. A. Rec. Ind. Mus., vol. 14, p. 25, Plate 8, fig. 4.

1928 Ventidiiis distanti Esaki, T. Ann. Mag. Nat. Hist. ser. 10, vol. 2, p. 511.

1929 Ventidiiis distanti Dover, C. Treubia, Deel 10, Afl. 1, Biz. 69 (part. ?).

1930 Ventidiiis distanti Esaki, T. Jour. Fed. Malay Mus., vol. 16, p. 18 (consid-
ers it to be a good .species ) .

1933 Ventidiiis distanti Lundblad, O. Arch. Hydrobiol. 1933 Suppl. Bd. 12,
Tropische Binnengewasser 4, S. 372.

In a paper entitled "Aquatic Rhynchota from the Southern Shan
States" Paiva described this species "from several specimens in al-
cohol from the top of the gorge of the He-Ho River, Yawnghwe State
ca. 3.500 ft., 7-iii-1917." We quote his description and reproduce
his illustration on plate 2.

"Apterous form. Head black with a large patch at base and a
transverse fascia at apex of face yellowish ochraceous; eyes silvery
grey, with a black patch on the disk; antennae black, basal half of
first joint yellowish.

"Pronotum very short, black, a narrow ochraceous waved fascia
at basal margin, anterior margin slightly concave, posterior margin
almost straight. Mesonotum large, about as long as its greatest
breadth, covered with decumbent hairs, disk obliquely striate on
anterior area, ochraceous, with two broad lateral black fasciae curved
inwards anteriorly and meeting narrowly on anterior margin, each
extended posteriorly to meet a curved fascia on the intermediate
acetabula; a large subtriangular patch at centre of posterior margin;
the posterior lateral angles narrow!)- dull black.

Concerning the Genus Ventidius 333

"Metanotiim dull black with a small ochraceous spot near each
basal angle.

"Abdomen above dull white, the basal segment, a spot at lateral
margin of each segment and the apical segment black.

"Underside pale ochraceous; legs black, base of anterior femora
ochraceous.

"Length 3 mm."

Type No. 7125/ H. I. in the collection of the Zoological Survey of
India.

Dr. A. P. Kapur of the Calcutta Museum is now unable to find this
type in their museum and it may have been lost by high water in the
temporary quarters in 1943. This is most unfortunate since only a
re-examination of the type or a study of a series of specimens from
the type locality the identity of Ventidius distanti can be established.

Dover ( 1928 ) states that "Paiva's V. distanti is based on apterous
males which are very different in coloration from the females. I
have taken specimens in Kuala Lumpur which agree exactly with
Paiva's description and figure in copulation with specimens as de-
scribed by Distant." We know that Dover had two species before
him from Selangor, F. M. S. and had labelled the larger species as
"Metrocoris aquarius Distant" in error. This we have described as
a new species. The smaller species which he had from Selangor,
F. M. S. is at least subgenerically distinct from Ventidius and the
males are less than 3 mm. long and the pronotum is entirely black
(Paiva gives for V. distanti 3 mm. long and a narrow ochraceous
waved fascia at basal margin of the very short black pronotum).
Whether this can be V. distanti or not we cannot determine. Both
Esaki and China have determined these specimens as V. distanti.
Since, besides their smaller size and different color pattern, the males
have a tubercle on the ventral side of the front femur and a black
tubercle on the mesosternum that wre not mentioned in Paiva's de-
scription of V. distanti we must (juestion their determination. It
may belong to the new subgenus Ventidiuides along with V. kuiferti
sp. nov. or be something quite different.

Ventidius (Vcntidioides) kuiterii sp. nov.
(Text fig. 1, PI. 2, figs. 2, 3, Pi. 3, figs. 6, 7)
Size: Wingless male: 2.3 mm. to 2.5 mm. long; across metaace-
tabula 1.57 mm. to 1.59 mm. wide. Wingless female: 2.7 mm. to
2.95 mm. long; across metaacetabula 2.1 mm. wide.

Color: Black with whitish or yellowish-white figures as shown
on plate. These figures somewhat variable in shape and size.

334

The University Science Bulletin

Metanotal pale spots sometimes joining the posterior lobes of
mesonotal pale spot. Abdominal spots sometimes joining posterior
lobes of mesonotal pale spot. Abdominal spots vary from three to
seven in number. Connexival segments of female whitish, of males
black except last two. Metacetabiilar elongate whitish spot with
bidentate lower margin. Venter and coxae whitish. Antennae
black except base of first segment and third segment sometimes
pale. Legs black except base of front femora.

Structural charoct eristics: Antennae as long as body in male,
somewhat shorter than body in female. Relative lengths of anten-
nal segments in male: 1st: 2nd: 3rd: 4th: :60: 18:20:24; and those in
female :: 47: 15: 23: 22. Eyes in wingless forms overlapping lateral
margins of prothorax and anterolateral angles of mesothorax. An-
terior and posterior margins of pronotum medially concave. Meso-
notum swollen. Metanotum more or less declivous. Interseg-
mental suture between meso- and metanota as well as anterior
margin of first tergite obliterated or lost. Abdomen reduced.

Relative lengths of leg segments of a male.

Femur

Tibia

First
tarsal

Second
tarsal

Total

of
tarsus

Front leg

140
388
430

115
205
140

110

34
26
30

Middle leg

Hind leg

136
54

Connexivum relatively broad. Basal abdominal tergites more or
less fused, only the last five distinct. Last ventral abdominal seg-
ment as long as five preceding segments together in both sexes.
Front femur slightly curved in basal third and that of male some-
what thickened and with a tubercle on the middle of ventral side.
Right paramere of male smaller and somewhat different in shape
from the left paramere ( see pi. 3 figs. 6, 7 ) . The male also possesses
a median dark tipped tubercle* on the mesosternum that appears
to be the caudal end of a tube. This is far in front of the omphalium
which lies on the small triangular metasternal piece. Females lack
this tubercle and males of other species of Ventidius must lack it
since it has not been described (see pi. 3 figs. 6, 7).

* We cleared a male in caustic potash and mounted the thoracic venter on a slide.
Under 16 mm. objective of the compound microscope (lOOx) the dark spot on the tip of
the tubercle is at the caudal end of a long median longitudinal duct. Under 4 mm. ob-
jective ( 440x ) the dark spot is a cluster of minute setae but no definite pore-opening for
the duct is visible. The setae resemble glandular setae arising from circular trichophores.

Concerning the Genus Ventidius

335

Location of types: Described from fifteen males and thirty-five
females bearing the label "Shingbwiyang, Burma III 24-1944 Lt.
L. C. Kuitert," and five males and six females from the same area
on VIII, 17-1944 by L. C. Kuitert. The male holotypes, female

V. kuiterti

Text fig. 1. Ventidius (Ventidioides) kuiterti sp. no\ .

A. Ventral view.

B. The tubercle enlarged (X 440).

C. The male front leg.

D. The male antenna.

336 The University Science Bulletin

allotype and paratypes are in the Francis Huntington Snow Mu-
seum, University of Kansas.

Comparative notes: This species is the first one described in the
genus Vent id ins that shows sexual dimorphism in the front leg and
the mesosternum. The males have a vential projection on the front
femur and a mesosternal tubercle.

Separation of Ventidius Distant from Esakia Lundhlad and Key to the

Species of Ventidius

1. In wingless forms lateral longitudinal suture of metanotum reach-
ing intersegmental suture between meso- and metanota. Third and
fourth segments of male antennae somewhat enlarged and fringed
with hairs Esakia Lundblad

1'. In wingless forms lateral longitudinal suture of metanotum not
reaching intersegmental suture between meso and metanota. Third
and fourth segment of male antennae normal. . . Ventidius Distant (2)

2. Posterolateral angle of metaacetabula transverse and broad or
oblique. Males without a tubercle on front femur, and without

a small tubercle on mesosternum Ventidius (Ventidius) ( 3 )

2'. Posterolateral angle of metaacetabula bilobate. Males with a tu-
bercle on front femur and with a small black tubercle on meso-
sternum. Parameres of male asymmetrical and large.

Ventidius (Ventidioides) subgen. nov. (9)

3. Front tarsal claws arising from near base of second tarsal segment
and male with a tuft of hairs on third antennal segment.

V. (Ventidius) malmjensis sp. nov.
3'. Front tarsal claws arising from nearer the middle than base of

second segment. Male antennae without tuft of hairs (4)

4. Size rather large (3.5 mm. long or longer) (5)

4'. Size smaller (less than 3 mm. long) (6)

5. Antennal segments three and four about equal in length. Winged
form with a T-shaped black figure on pronotum.

V. (Ventidius) uquarius Distant

5'. Antennal segment three considerably longer than four (38:28).

Winged form with a median longitudinal pale line on pronotum.

V. (Ventidius) usingeri sp. nov.

6. In wingless forms intersegmental suture between meso- and meta-
nota distinct; intersegmental suture between metanotum and first
tergite especially distinct. Parameres of male club shaped with
a protuberance on rear margin V. ( Ventidius) modulatus Lundblad

7. Upper surface of body with considerably large brown or black

areas, head with brown or black spots (8)

7'. Upper surface of body largely stramineous, head entirely stramine-
ous V. (Ventidius) chinai sp. nov.

8. Pronotum black with a pale transverse triangular area.

V. (Ventidius) werneri sp. nov.

Concerning the Genus Ventidius 337

8'. Pronotum solid black or brown except on winged form whicb has

two pale spots V. (Ventidius) Jicnryi Esaki

9. Stramineous metanotal lobes very broadly continuous to meso-
notal spot. Base of pronotum transversely white.

V. (Ventidioides) distanti Paiva (?)*
9'. Stramineous metanotal lobes not or narrowly continuous to meso-
notal spot. Pronotum entirely black.

V. (Ventidioides) kuiterti sp. nov.

* Based on determinations by Drs. China and Esaki.

338 The University Science Bulletin

PLATE I

1. V. (Ventidius) aquarius Distant. Winged form (copied from Distant,
1910).

2. V. (Ventidius) aquarius Distant. Wingless female (?) (copied from
Distant).

3. V. (Ventidius) henryi Esaki. Winged female.

4. V. (Ventidius) henryi Esaki. Wingless male.

5. V. (Ventidius) chinai sp. nov. Wingless male.

6. V. (Ventidius) chinai sp. nov. Wingless female.

7. V. (Ventidius) modulatus Lundblad. Wingless male (copied from
Lundblad, 1933).

8. V^. (Ventidius) malayensis sp. nov. Wingless female.

Concerning the Genus Ventidius

339

PLATE I

V. aquarius

V. henry

V.chinai

6 V.chinai 7 V modulatus

8 V. HDobyensis

340 The University Science Bulletin

PLATE II

1. V. (Ventidiits) disianti Paiva. Wingless female (copied from Paiva
1918).

2. V. (Ventidius) kuiterti sp. nov. Wingless male.

3. V. (Ventidioides) kuiterti sp. nov. Wingless female.

4. V. (Ventidius) wemeri sp. nov. Wingless male.

5. V. (Ventidius) wemeri sp. nov. Wingless female.

6. V. (Ventidius) henryi Esaki. Wingless female.

7. V. (Ventidius) usingeri sp. nov. Winged female.

8. V. (Ventidius) usingeri sp. nov. ^^'ingless female.

Concerning the Genus Ventidius

341

PLATE II

V. distanfi

5
V. werneri

6 V.henryi

8
V. usingeri

342 The University Science Bulletin

PLATE III

1. E.sakia usingeri sp. nov.

2. A hypothetically primitive species of VenticUus with all thoracic and
abdominal sutures retained.

3. The male antenna of V. (Ventiditts) maJmjcnsis sp. nov.

4. The male front leg of V. (Ventiditis) usin^eri sp. nov.

5. The male front leg of V. (Ventiditts) malayensis sp. nov.

6. The left paramere of V. (Ventidioides) kuiterti sp. nox .

7. The right paramere of V. (Ventidioides) kuiterti sp. nov. i

8. The left paramere of V. (Ventiditts) henrtji sp. nov. \

9. The left paramere of V. (Ventiditts) chinai sp. nov. \

10. The left paramere of V. (Ventidius) usingeri sp. nov. '

11. The right paramere of V. (Ventiditts) tcerneri sp. nov.

12. The left paramere of V. (Ventiditts) malatjensis sp. nov.

Concerning the Genus Ventidius

343

PLATE III

^MESONOTUM^-^

^INTEBSEGMENTAL

SUTURE jy^^^

^-. METANOTUM--tf

t LATERAL SUTURE'

-.1ST ^BD y

SPIRACLE^^ ^ ^
1ST ABD. SEGMENT^ 2

Esakia usinqeri

Hypothetically prlmilive
Ventidius species

■^===::^>

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XL] April 20, 1960 [No. 8

A Revision of the Mite Family Bdellidae in North and
Central America (Acarina, Prostigmata)

BY
Warren T. Atyeo

Abstract. This study, although primarily a monographic revision, includes
data on the bionomics, morphology, and intraspecific variation of the bdcllids.
Eleven genera are recognized, including a new genus Octohdellodes; four gen-
era or subgenera are synonymized, these are: Caenohdella Oudemans, 1937
(=: Bdella), Troglobdella Oudemans, 1937 (^Cyta), Hoplomolgtis Bcrlese,
1923 (= Neomolgus), and Hoploscirus Thor, 1937 {= Bdellodes).

Thirty-two species are included in this paper, of which sixteen are new.
Four species are moved to other genera, and thirteen species, for which type'
materials arc available, are s\nonymized. For other synonymies, Thor ( 1931 )
is recognized.

The sixteen new species described are: Bdella longistriata, B. tropica,
Bdellodes hisetosa, Cyta spuria, Neomolgus rnutahilis, Octohdellodes hurdi,
O. infrequens, Odontoscinis iota. O. alphinus, Thoribdella communis, T. insolita,
T. simplex, T. spiuosa, T. communis, Spinihdella bifurcatci, S. ornata.

TABLE OF CONTENTS

Introduction 346

Acknowledgments 346

Collection of Material =. 347

Preparation of Material 347

Geographical Distribution 348

Bionomics 349

Morphology 351

General 351

Segmentation of Appendages 351

Setae 352

Chaetotaxy of the Legs 353

Gnathosoma 355

Propodosoma 357

Dorsal I l>ster<)soma 358

Anal Hcgion 359

Genital Region 359

(345)

346 The University Science Bulletin

PAGE

Systematic Relationships 361

Intraspecific Variation 365

Taxonomy 365

Historical Account 365

Characters and Descriptive Methods 367

Description of the Family Bdelliuae 370

Key to the Subfamilies and Genera of the Bdellidae 37 1

Subfamily Bdellinae 371

SubtiuniK- Otlontoscirinae 385

Subfamily Cytinae 416

Subtamih' Spinibdellinae 423

Selected Bibliography' 439

INTRODUCTION

The purpose of this study has been to develop a solid basis for
the systematics of the family Bdellidae. Inadecjuate descriptions,
loss of types, and conflicting impressions of the type genus BdcIIa
often make it wholly impossible to determine which of the previously
proposed species are valid.

Of the twenty-five hundred specimens examined, approximately
twenty-one hundred were from Central and North America, two
hundred from Iceland, one hundred from Australia, one hundred
from Europe, and less than five from each of the continents South
America, Asia, and Africa. Although over one hundred species and
eighteen varieties have been described in the literature, only twenty
named species were discovered, of which, sixteen occmred in Cen-
tral and North America. Thor ( 1931 ) synonymized many species
with those that appeared to him to be valid. Thor's synonymies
will not be (piestioned in the present investigation, which, due to the
lack of abundant European material, is necessarily restricted in
scope.

To clarify my position in the taxonomy and systematics of this
group, detailed descriptions and illustrations have been introduced
for the known species of Central and North America, which number
sixteen named and sixteen new species.

ACKNOWLEDGMENTS

To Robert E. Beer and Charles D. Michener, of the University
of Kansas, I am indebted for guidance and invaluable suggestions
during the course of this study. Further appreciation is acknowl-
edged to Joseph H. Camin of the University of Kansas for assistance
in the preparation of portions of this manuscript and to Edward W.

A Revision of the Mite Family Bdellidae 347

Baker of the United States Department of Agriculture for the loan
of type materials. Thanks are due to S. L. Tuxen of the Universi-
tetets Zoologiske Museum, Copenhagen; K. Strenzke of the Max-
Planck-Institut fur Meeresbiologie, Wilhelmshaven; D. M. Allred
of the University of Utah; and to the many other workers who have
generously contributed specimens for this study.

COLLECTION OF MATERIAL

It is desirable to collect and preserve these mites in such a way
as to avoid contamination with fine particles of dirt and sand. Both
of these substances readily adhere to the integument and, even with
drastic clearing methods, so much accumulation may remain on
the specimen that even generic determinations are difficult.

Berlese funnels efficiently extract bdellids from litter, soil, nests,
and compact jilants, such as mosses and foliose lichens. Water is
superior to alcohol in the Berlese collecting vial. The high surface
tension of water, as compared to alcohol, will support many of the
mites on the surface while permitting soil particles to sink. With a
small brush or spatula, the mites can be transferred to a preservative
with a minimum of extraneous matter.

To collect from moss, lichens, and other miscellaneous epiphytes.
en situ, one can beat the material over a white enamel pan and re-
move the dislodged mites from the pan with a small brush. Sim-
ilarly, individual mites can be removed from rocks, boards and other
relatively large and smooth surfaces.

Habitat specificity has not been demonstrated in the Bdellidae,
although this may be due to a lack of data that would mask the
character of the true situation in this regard.

PREPARATION OF MATERIAL

Two satisfactory media for mounting bdellids, which are soft-
bodied mites, have been used. These are polyvinyl alcohol with
lactic acid and phenol added (PVA-L-P) and Hoyer's modification
of Berlese's mounting medium (see Beer, 1954, for formulations).
Comparing these two media, each has certain advantages and dis-
advantages. PVA-L-P has less clearing action and it is less difficult
to prepare specimens in which appendages are in the same hori-
zontal plane, but to remount from this medium is an extremely haz-
ardous undertaking with specimens having a soft integument.
Hoyer's clearing properties are excellent for smaller uncleared bdel-
lids, causing little distortion in striations, shape or size. Although

348 The University Science Bulletin

slightl)- more diffiiciilt to use than PVA-L-P, the ease with which
specimens can be remounted without the loss of or damage to parts
is an advantage that cannot be over-emphasized.

Characters necessary for specific determinations are located on
both the dorsal and ventral smfaces of the body. For this reason,
large and opaque specimens must have their body contents removed
or thoroughly cleared before mounting. Other specific characters
are found on the surfaces of the legs and palps. For easy study, it
is therefore desirable to have these appendages similarly oriented
and extended in the same horizontal plane. Heating of specimens
in lacto-phenol at 200' to 250^ F. for a short time will soften the leg
muscles sufficiently to allow easy manipulation of these appendages
and further heating,in lacto-phenol slightly under the boiling point
(up to 400' F. after moderate evaporation) will dissolve the body
contents. The fime required for the latter process is variable, de-
pending on the size of the specimen and the type of preservative.
Potassium' hydroxide, although used as a clearing agent in many
groups, is not a satisfactory reagent for bdellids as its drastic action
causes a considerable loss of diagnostic setae.

Specimens used for the study of musculature were prepared by
a different process. Bdellids, previously preserved in ninety-five
per cent alcohol, were cleared in beechwood creosote for five to ten
minutes, then transferred directly to thin balsam. The palps and
legs were removed from the body with small needles and placed in
the mounting medium. A cover slip, first immersed in xylene, was
gently pressed against the specimen until the appendages were in the
same horizontal plane.

GEOGBAPHICAL DISTRIBUTION

Mites of the family Bdellidae have been collected from all major
land masses of the world and from many insular groups. When dis-
tribution records are more complete, many species will probably be
found to be cosmopolitan in distribution. Many of the species found
in the Western Hemisphere are also found in other parts of the
world. One species is known to occur in North America from
Mexico to Alaska, as well as in Iceland and Europe. One species
has been collected in China, Australia, Hawaii, and North America.
Another species is circumpolar in distribution.

A Revision of the Mite Family Bdellidae 349

BIONOMICS

Bdellids are active, fast-running mites, predaceous on small arthro-
pods and arthropod eggs. They seem to occur in almost every ter-
restrial habitat where food material is available. Three investiga-
tions on the biology of these mites have been reported. Womersley
(1933a) and Currie (1933) studied one species as a possible bio-
logical control agent for the clover springtail or lucerne flea (Smin-
thunis viridis L.) and Snetsinger (1956) studied the biology of
S])iiiibdella depressa (Ewing).

Species observed in temperate regions apparently have broad
temperature and humidity tolerances. Although specimens are
most frequently collected in moist, cool habitats, this may be a
consequence of the availability of food animals. Many individuals
have been collected from dry, exposed surfaces, and Bornemissza
(in litt.) has evidence indicating that a Cyfa species may be re-
stricted to the desert regions in Australia. On the other hand, Snet-
singer (1956) could obtain complete life histories only at ninety per
cent relative humidity between sixty and seventy degrees Fahrenheit.

Bdellids in temperate climates overwinter in all stages of develop-
ment. In Kansas, the author has collected all the active stages of
local Cyta and Bdclla species from leaf litter as early as February
22, and although the deutonymphal and tritouymphal stages were
the most prevalent, the larvae, protonymphs, and adults were also
present. Snetsinger ( 1956 ) observed that in Illinois the egg stage
is the most abundant overwintering form of Spinihdella depressa,
but all stages except the larval stage were found hibernating under
tree bark.

These mites have the life history stages found in many trom-
bidiform mites, that is, egg, deutovum, larva, three nymphal stages,
and adult. Each active immature stage ends in a period of
quiesence, following which the old integument is shed. Currie
(1933) reported that one nymph of Biscirus Japidarius consumed
eighteen immature clover springtails on each of three successive
days, and Snetsinger (1956) reported that immatures of SpinihdcUa
depressa require three or more tetranychid mites to complete each
developmental stage. The latter author also found that the length of
time necessary for development is partially dependent on tempera-
ture, lengthening as the temperature decreases. Development from
larva to adult at ninety percent relative humidity required twenty-
one to thirty days at sixty degrees Fahrenheit, but only fourteen to
twenty-one days at seventy degrees.

350 The University Science Bulletin

Oviposition preferenda have not been reported; however, there
is a tendency for the females to lay eggs in protected areas. In
stender dishes coated with a plaster-of-paris and charcoal mixture
( Lipovsky, 1953 ) and marked with a deep line to simulate a crack,
the author observed the sites of egg deposition of four females.
Sixteen spiny elliptical eggs, laid singly at the rate of one egg each
one and one-half days, were deposited in the prepared crevice at
the bottom of the dish, while only one egg was laid in an exposed
area near the wall of the dish. In the field, Snetsinger ( 1956 )
found that large numbers of eggs were deposited during the autumn
months under the basal bark of trees, probably a result of females
aggregating in protected spots at the onset of cold weather.

Parthenogenesis has not been discovered in the Bdellidae. Al-
though the forty-five specimens of Bclella tropica, sp. nov. and one
hundred and forty-six specimens of SpinihdcUa cronini examined
in this study were all females, this cannot be construed as proof of
parthenogenetic development. In all species for which large num-
bers of specimens were available, there were usually many more
females than males. For example, there were one hundred and
fifty-six females and forty males of Bdella lon^icornis and ninety-
two females and only four males of Bdella niiisconini. The sexes
were approximately equal in number only in Bdellodes longiroslris,
of which forty-eight females and forty-four males were examined.
Occasionally in small collections of two to ten specimens of the
three latter species, all or most of the individuals were males.
Plausible explanations for the scarcity of males, especially for
B. tropica and S. cronini, could be that the sex ratio is extremely
unequal; that the males have different habitat preferences; or that
they have different peaks of seasonal abundance.

Personal obserxations on feeding habits were made by the use
of covered stender dishes with collembola as the food source.
Bdellids, when placed with collembola, move slowly until contact
is made with the distal setae of the palpi. If the mite is startled,
it will run rapidly backwards. If not startled, the mite will lunge
at the prey, impaling it on the tips of the mouth parts. The chelic-
erae are rapidly and alternately extended and retracted at various
angles into the body of the prey, while the chelae are opened and
closed, thus macerating the tissues. If disturbed while feeding, the
bdellid elevates the tip of the gnathosoma with the mouth parts still
inserted in the prey, and in this fashion the mite may move away
from distracting influences. When feeding is completed, the mite
either goes to a secluded spot or again moves slowly about the dish.

A Revision of the Mite Family Bdellidae 351

MORPHOLOGY
General

The general body configuration of the bdellids has been known
for many years. Geoffroy's (1762) drawing of Bdella longicornis,
even with its inaccuracies, can easily be recognized as belonging to
this family. The knowledge of the external morphology has in-
creased with better optical equipment, culminating with Grand-
jean's (1938) investigation. The internal morphology is known
only from Michael's (1896) study of the internal anatomy of
Neomolgus liltoraUs {=^ Bdella hasteri Johnston, 1836).

The body is divided into three distinct regions, the gnathosoma,
propodosoma. and hysterosoma. The anterior gnathosoma con-
sists of two elongate, chelate chelicerae (mandibles of authors), a
ventral hypostome (rostrum of authors), and two geniculate palpi.
Collectively the chelicerae and hypostome form a conelike structure
projecting from between the palpal bases. The propodosoma bears
the two anterior pairs of legs, eyes, and pseudostigmatic organs, this
region being delimited dorsoposteriorly by a conspicuous con-
striction. The hysterosoma bears the posterior legs, anus, and
genitalia.

Segmentation of Appendages

In order to determine the possible homologies between the legs
and the palpi, it is necessary to establish basic relationships in the
more primitive of the two types of appendages, the legs. Bdellid
larvae have seven-segmented legs (including the pretarsus), lack-
ing the division between the basifemur and the telofemur. The
division of the femur is barely perceptible in the anterior three pairs
of legs in the protonymph; but each succeeding instar exhibits a
progressively more-distinct femoral division in all legs. Gamin has
prepared diagrammatic illustrations of the musculature of the legs
and palpi (figs. 3-5). According to Gamin's forthcoming theory
(in manuscript), he believes that, primitively, each primary seg-
ment of the leg had two flexor muscles which insert on the succeed-
ing segment. In bdellids, the three distal segments have undergone
considerable change and may not represent the primitive genu,
tibia, and tarsus, but for convenience they will be denoted by the
standard terminology in this paper. The primitive condition is best
demonstrated in the femur which has two flexor muscles (fig. 3,
a, a) inserted at the base of the genu. An advanced feature, the
ventral flexor muscle of the telofemur may be a branch of the proxi-

352 The Universit\' Science Bulletin

mal flexor of the genu, the insertion of which has migrated. The
undivided femur in the larva and the lack of muscles inserting on
the telofemur in all developmental stages suggest that the telofemur
is not a primary segment but a secondary subdivision of the femur
allowing additional flexibility of the legs.

Previous workers have referred to the three distal segments of
the palpus as the genu, tibia, and tarsus. Although the musculature
of the palpus is greatly reduced, it is possible to homologize the
palpal segments with the segments of the leg, thus demonstrating
that these segments represent the telofemur, genu, and fused tibio-
tarsus. The femur, partially divided into basi- and telofemur, has
two long flexor muscles ( fig. 5, o, a ) homologous to flexor muscles in
the legs; the small extensor muscle (fig. 5, /) has no homologue in
the legs. Two muscles inserting at the base of the distal segment,
with origins in the penultimate segment and telofemur, can be
homologized with the flexors of the tibiae of the legs. The penulti-
mate segment of the palpus can therefore be homologized with the
genu of the leg. The distal palpal segment, lacking muscles and
pretarsal elements, represents the remaining segments, the tibia,
tarsus, and pretarsus. In the Cunaxidae, the musculature of the
palpus (fig. 6) is similar to that in the bdellids, but there is an
incomplete fusion of the tibia and tarsus.

Setae

Two general types of setae are abundant: thick-walled (tactile)
setae and thin-walled (chemosensory) setae. The tactile setae
("Eigentlichen Haare" of Vitzthum, "Foils proprement dits" of
Grandjean) are inserted in alveoli, are movable, tapering, nude
or plumose, and have extremely small lumina. The chemosensory
setae ( "solenidions" of Grandjean ) , for convenience termed sensory
setae, are inserted in small pits or depressions lacking alveoli; they
are fixed, nude or minutely pilose, and have large lumina. Com-
monly, the form of the thin-walled setae is slender and tapering
(attenuate) (fig. 189), or thick and broadly rounded distally (fig.
188). The tips of both forms curve away from the body. In bdel-
lids, the sensory setae are not striated as in many other groups of
prostigmatic mites. Sensory setae are restricted to the four distal
segments of the legs except for one on the palpal tibiotarsus.

The long apical setae of the palpal tibiotarsus, which are tactile
in function, have two types of insertions. In Cyta and a few species
of Bdella and SpinihdeUa, the setae originate deep within the seg-

A Revision of the Mite Family Bdellidae 353

ment; in the other genera studied, the apical setae are sHghtly en-
larged at their bases and inserted in superficial sockets. Commonly
in the Odontoscirinae, the edges of the apical sockets are heavily
sclerotized and form distinct rims.

The four sensilla, although of variable lengths and diameters, are
long and tapering (except Thoribdella) and each is inserted in a
pseudostigmatic organ. In the genus Thorihdello, the posterior
sensilla vary from short thin setae to short thickened rods which
are divided in the middle into three connecting arms (fig. 150).

A third type of seta is short and peglike and has a variety of
shapes and forms (figs. 190-193). Solid pegs, possibly blunt spines,
are found on the dorsolateral rims of coxae I and II (fig. 2, /) and
on the dorsolateral regions of the hypostomal bases in close prox-
imity to the cheliceral bases ("L'cpine laterocoxale du palpe" of
Grandjean). The positions of these three pairs of setae suggest
that they function as proprioceptors. Other pegs, maximally one
on each of the two distal segments of legs I and II, are inserted in
pits, lack alveoli, and may represent highly modified chemosensory
setae.

Chaetotaxy of the Legs

The chaetotaxy of the legs provides important taxonomic char-
acters. The majority of the leg setae are tactile, and sensory setae
and pegs occur only on the dorsal surfaces. Tactile setae are ar-
ranged on the legs as follows: two or more rows of ventral setae
that tend to be plumose on the tarsi; one row each on the anterior
and posterior (lateral) surfaces, and one to three unpaired setae
on the dorsal surface of each segment except the coxa.

The majority of the sensory setae occur on legs I and II, with leg

I having the greater number. Tarsus I has two, occasionally three,
broadly rounded sensory setae (fig. 188), two or more attenuate
sensory setae (fig. 189), and usually one peg (figs. 190-193). Tarsus

II usually has the same number of blunt sensory setae and pegs, but
fewer attenuate sensory setae than tarsus I. The proximal halves
of tibiae I and II have only attenuate sensory setae, but the distal
portions may have any combination of the various types of special-
ized setae. The tibiae and tarsi of legs III and I\' have only the
attenuate type of sensory setae.

Duplex setae similar to those found in the Tetranychidae (Pricli-
ard and Baker, 1955) and/or attenuate scnsor\' setae are inserted on
the genua. When duplex setae are present, they occur on the
anterior three genua, but may be wanting on genu IV. The inser-

12—8920

354 The University Science Bulletin

tions of the micro- and macrosetae that comprise a duplex seta may
be confluent, separate and approximate, or separate and distant
(figs. 194-197).

The number of tactile setae may vary considerably within a
species, as is the case with the attenuate sensory setae on the genua
and proximal halves of tibiae I and II. The other types of sensory
setae are usually constant in number, but may vary in position ( see
p. 365).

Trichoboths (long tactile setae) are inserted in deep, heavily
sclerotized sockets (fig. 184) and are similar in structure to the
dorsal sensilla. The maximum number of trichoboths is five pairs,
which are inserted on tibiae I, II, and IV and tarsi III and IV. The
subfamilies are characterized, in part, by the number of trichoboths
present in the adult stage. The Odontoscirinae have five pairs ar-
ranged as above; the Spinibdellinae and Bdellinae have four pairs,
lacking the trichoboth on tibia II (Bdella mexicana also lacks the
trichoboth on tarsus IV); and the Cytinae have less than four pairs.
In the latter subfamily, trichoboths are inserted on tibiae I and IV
and tarsus III except Cijta latirostris which has only one trichoboth
inserted on tibia IV. In species with less than five pairs of tricho-
boths there is a large tactile seta in the position normally occupied
by a trichoboth when compared with the Odontoscirinae.

The larvae of all species studied (except Cijta latirostris) have one
trichoboth which is inserted on tarsus III; the protonymphs have
the same number of trichoboths as the adult on the anterior three
pairs of legs, but leg IV has none; succeeding instars have the adult
number on all legs. C latirostris is an exception, having one tricho-
both inserted on tibia IV in the tritonymphal and adult stages, the
other instars lack trichoboths.

The extremities of all the tarsi are abruptly narrowed subapically
where the dorsal tarsal surfaces bend sharply downward to the pre-
tarsus. On each of the oblique surfaces thus formed are three pairs
of setae which constitute the dorsoterminals. The proximal pair of
dorsoterminals {dt 1) are long and curve obliquely from the claws
(leg IV may have only one seta in this position). The second pair
{dt 2) is inserted midway between dt 1 and the pretarsus, and the
distal pair (dt 3) flanks the pretarsus. The two distal pairs (dt 2, 3)
extend to the bases of claws and are present in all known species ex-
cept Biscirus silvaticiis which lacks one of the middle pairs of setae.
The dorsoterminals may be thick-walled (termed solid) or thin-
walled (termed hollow) and may be nude to coarsely branched

A Revision of the Mite Family Bdellidae 355

(figs. 179-183, 185-187). Although different forms may occur on
different legs, the structure of the dorsoterminals is usually constant
within a species. Generally, thin-walled setae on the anterior legs
are replaced by thick-walled setae on the posterior legs.

The short pretarsus, indistinctly divided on its dorsal surface into
three pseudosegments, arises between dt 3. The distally inserted
claws may have one, two, or no small dorsal ridges bearing laterally
directed rays (fig. 70). These rays may be long and few in number
or minute and numerous. In the latter case, the small rays are des-
ignated as the minute rays. Only the size of the claws and the pres-
ence or absence of lateral and/or minute rays were found to be of
taxonomic value.

Gnathosoma

The gnathosoma, consisting of the elongate chelicerae, hypostome
and palpi, is one of the most characteristic features of the Bdellidae.
The geniculate palpus has six segments: coxa, trochanter, basifemur,
telofemur, genu and tibiotarsus, the latter bearing apically one or
two long tactile setae. The coxa is fused with the base of the hypo-
stome, and thus is indistinguishable as a separate segment. The tro-
chanter is small and devoid of setae. The femur is incompletely di-
vided into a long proximal basifemur and a short distal telofemur.
The basifemur bears a variable number of setae, but the telofemur
has only one seta that is always inserted on the dorsal surface. The
genu may be as long or longer than the telofemur and bears two to
seven setae. The terminal, fused tibiotarsus varies greatly in length
and shape. Typically, the tibiotarsus is approximately the length
of the genu plus the telofemur, is expanded distally, and obliquely
truncated at the apex; or it may be more than twice as long as the
genu plus the telofemur, cylindrical, and apically rounded. Un-
fortunately, there are many intergrades between these two readily
recognized conditions, thereby making size and shape of the palpal
tibiotarsus a highly subjective character in many instances.

The dorsal and ventral apical setae resemble the trichoboths of
the legs and the dorsal sensilla. In the Odontoscirinae, the end setae
are approximately equal in length and shorter than the cylindrical
tibiotarsus. In MonotrichohcJeUa, there is a single apical seta. In
the remaining genera, the end setae are subequal and usually much
longer than the tibiotarsus.

In life, the geniculate palpi are flexed at the two distal articula-
tions and are carried extending forward and bowed upward. In
slide preparations, especially with cleared specimens, the medial or

356 The University Science Bulletin

lateral aspects are most frequently encountered. Two useful points
for determining orientation of the palpus are the relative positions
of the dorsal and ventral end setae and the position of the single
dorsal seta of the telofemur.

The chelicerae are elongated and bear movable chelae. The
number and position of setae, the shape of the chelicerae, and the
form of the chelae are sufficient to distinguish many genera.

The majority of the genera have normal chelicerae which are
approximately three to five times as wide at their thickest portion
as at their thinnest, and have chelae with small sickle-shaped mov-
able digits and small straight fixed digits (figs. 7-8). The inflated
chelicerae of a few species of Odontoscirus and BdcUa (figs. 13, 16)
are more than five times wider at the base than at their narrowest
parts. In Odontoscirus the inner surface of the movable digit is
straight if the teeth are disregarded (figs. 9, 16, 17). A third chelic-
eral shape occurs in the Spinibdellinae (figs. 32-35) which has a
narrow form with almost parallel sides and needlelike chelae (fig.
10 ) . The genus Cijta is unique in having large, thickened chelicerae
(figs. 30-31) tipped with massive chelae (fig. 11). One or more
teeth may be present on the inner surfaces of the fixed digits.

Two setae are inserted on the dorsal or dorsolateral surfaces of
the chelicerae in most genera. Cyta has two setae on each chelicera,
but one is inserted at the base of the fixed digit (fig. 11 ); Neomolgus
has more than two setae; and Bdellodes longirostris and one unde-
scribed species of Thoribdella from Australia have only one seta on
each chelicera.

Viewed from the ventral aspect, the gnathosoma has three dis-
tinct regions ( fig. 2 ) : a transversely striated, rectangular base
bearing the palpi at the anterolateral angles; a long, gently ta-
pering buccal cone; and two small, terminally fringed lateral lips
which are separated from the buccal cone by a weak suture. The
rectangular gnathosomal base consists of the palpal coxae fused with
a basal portion of the hypostome, and the buccal cone and the
lateral lips represent the major region of the hypostome.

The setae inserted on the gnathosoma can be best characterized
by their positions. The ventral hypostomals {vh series) are on the
ventrolateral surfaces of the buccal cone and the anterior margin of
the hypostomal base. In adults, either two, six, or seven pairs of
these setae are present. Two pairs of small adornal setae inserted
on the lateral lips are present in all the developmental stages.
Finally, one pair of setae may be present on the dorsal surface of

A Revision of the Mite Family Bdellidae 357

the hypostome; when present, these setae are covered by the bases
of the cheHcerae ("les polls premandibulaires" of Grandjean).
When characteristic for a species, these dorsal hypostomal setae
occur in all the stages of development.

In genera with six or seven pairs of ventral hypostomal se'ae, the
immature stages may be identified by the number of setae present.
The larvae have two pairs; the deutonymphs, four pairs; and the
tritonymphs have five pairs. The Cytinae and Spinibdellinae have
two pairs of ventral hypostomal setae in all stages.

The dorsum of the propodosoma has a thicker integument than
the remainder of the idiosoma. A uniform thickening on the inner
surface has developed which is approximately rectangular to reni-
form in shape, and encompasses the four pseudostigmatic organs
("shield" of authors). This thickening of the integument at-
tains greatest development in the Odontoscirinae.

Internal ridges or apodemes connecting the pseudostigmatic
organs laterally and sometimes anteriorly have developed secondar-
ily in a few species. In their simplest form, the lateral apodemes
are narrow with parallel margins (fig. 140). More complex forms,
as in Bdella longicornis, have smaller thickenings mesal or lateral
to the prominent lateral apodemes which form an interlacing net-
work of supporting structures (figs. 133-135).

Two pairs of sensilla are inserted in the pseudostigmatic organs.
In all the genera except TJiorihdella, the pseudostigmata are cup-
like sockets lined with concentric rings of small ridges and often
are heavily sclerotized. In Thoribdella, the posterior pseudostig-
mata (figs. 142-149) are deep, goblet-shaped cavities of thin in-
tegument, lined with finely divided ridges which are perpendicular
to the insertions of the sensilla.

A pair of median propodosomak setae (fig. 1) is always present
between the posterior sensilla. In structure, these setae resemble
the dorsal hysterosomal setae. Another pair of setae, the lateral
propodosomals, may or may not be present. If present, they are
inserted lateral to a line connecting the lateral sensilla. In struc-
ture, the lateral propodosomals are usually simpler than the median
propodosomal setae.

The integumental striae between the dorsal sensilla form distinc-
tive shield patterns. A particular pattern may be unique to a
species, or as in the Odontoscirinae, a pattern may be common to
many species. Each striation is a sharp ridgelike external thicken-
ing of the integument with small breaks along the crest. In micro-

358 The University Science Bulletin

scopic examination, the breaks cause the striae to appear finely
broken (fig. 133), coarsely broken (fig. 156), or sparsely broken
(fig. 136).

The eyes are lateral or posterolateral to the posterior sensilla.
Two pairs are usually present, and the eyes of each pair may be
appro.ximate to distant. One species, Spinibdella depessa, has
the posterior eyes wanting, but their former positions are indicated
by teardrop patterns of the striae. In addition to the lateral eyes,
Cyta has a fifth eye between the anterior sensilla (fig. 165). A
fifth eye or protuberance in a similar position was reported by
Baker and Balock (1944) in their original description of Mono-
trichobdella maxoshurni, but this could not be distinguished in
the remounted type specimen.

The podocephalic canal (Grandjean, 1938) could not be ade-
quately examined in specimens mounted in Hoyer's or PVA-L-P.
Grandjean describes this canal as being an external groove in the
integument or an internal tube which originates on each side of
the dorsoposterior surface of the buccal cone, follows the contours
of the lateral body wall, and terminates above coxa I. Three tubu-
lar glands of unknown function open into the canal: one near the
laterocoxal spine of the palpus, one at the junction of the idiosoma
and the gnathosomal base, and one close to the termination of the
canal above coxa I.

Dorsal Hysterosoma

The dorsum of the hysterosoma has five transverse rows of setae
(fig. 1). Using the nomenclature of Oudemans {fide Vitzthum,
1943), these setae are:

1. internal and external humerals

2. internal dorsals

3. internal lumbrals

4. internal and external sacrals

5. internal and external clunals

The sacrals and clunals are arranged in gently to strongly curved,
transverse rows, or in extreme modifications, in a subrectangular
arrangement. In a few species, the external clunals are wanting.
The hysterosomal setae are nude to plumose and are usually constant
in form within a species. In a few species the form varies in
populations from different geographical areas. The longitudinal
intervals between the internal humerals and the internal dorsals,
the first interspaces, are used as a relative measurement of the
lengths of the internal humerals.

A Revision of the Mite Family Bdellidae 359

Anal Region

The posterior anal cleft is surrounded by striae (border striae)
which either parallel the entire length of the cleft or are parallel
to approximately the center of the cleft, then bend sharply laterad.
The setae of the anal region (figs. 1, 2) differ from the dorsal
hysterosomals in being shorter, thinner, and usually nude. One
or more pairs of setae situated in the region of the parallel stri-
ations are considered to be the anal setae, or anals. One pair of
setae near the dorsal termination of the anal cleft is termed the
posterior paranals, or postanals. Otlier setae, lateral to the anals
are the paranals.

Genital Region

Two longitudinally striated genital plates or flaps are situated on
the venter of the hysterosoma between the anal cleft and coxae IV.
Each plate bears a regular or irregular row of genital setae which
may vary slightly in position and number within a species. The area
surrounding the genital plates has bilaterally arranged pairs of para-
genital setae (figs. 2 g, 178 b). In the Cytinae, there is an unpaired
median seta immediately anterior to the genital plates (fig. 178 c).
Species of Spinibdella may have unpaired setae, but they occur
between coxae IV.

When the genitalia are in repose, a chamber or vestibule is formed
above the genital plates, the walls of which are contiguous with
the mesal edges of the plates. Three subequal genital discs, each
with a small spine of unknown function immediately caudad, are
on the lateral walls of the vestibule.

Because of the method of preparation, only sclerotized portions
of the genitalia remained for examination. The ovipositor is a large
membraneous tube which can be telescoped into the body or wholly
extruded. When in the latter position, the genital discs are car-
ried outside the body and are at the base of the ovipositor. Slightly
distal to the mid-length as many as four pairs of postmedial setae
may be present, and surrounding the tip of the ovipositor there are
six to ten pairs of subapical setae. A transverse cleft bisects the
apex, and two small, heavily sclerotized structures, probably glands,
are internal to the angles of this cleft.

The male genitalia are more complicated. In the pair of large
sclerotized plates, the amphioid sclerites (Michael, 1896), each has
the lateral edge bent dorsad to form a dorsal arm. Simple to highly
modified setae form an irregular series along the periphery of each

360 The University Science Bulletin

amphioid sclerite (figs. 169-177). The penis is a thin muscular
organ surrounded by a loose membranous sheath (Michael, 1896).
During copulation, the amphioid sclerites are everted, thus placing
the dorsal setae in a ventral position in contact with the exterior.
Evidently, the penis is connected to the dorsal arms of the sclerites
and carried outward with the eversion of these sclerites.

Specific differences in the male and female genitalia exist, but
only the males of Spinibdella and Cyta have differences striking
enough to warrant their use as a taxonomic character at this time.
Although males were present for only three species of Spinibdella,
the bizarre modifications of the amphioid setae appear to be specific
and stable characters. The setae of the three Cyta species lack
unique shapes, but have different arrangements and lengths. In
other genera, the setae lining the amphioid sclerites are similar in
structure, the slight differences in length and position could not be
evaluated. Differences in the female genitalia are slight and can
be detected only if the ovipositor is almost wholly extruded, a con-
dition of rare occurrence.

Genital tracheae, present in the Spinibdellinae and Cytinae, open
into the genital vestibule in front of the anterior pair of genital
discs (fig. 178 d). These paired structures are round and have
spiral thickenings which resemble the taenidia of insect tracheae
(except Trachymolgus, see below). The genital tracheae of the
Cytinae each have a single dichotomy near the origin, one branch
leading to the anterior tracheal opening near the cheliceral base,
the other branch ending above coxa I. In the Spinibdellinae. the
tracheae lack dichotomies; a single branch on each side of the body
ends above coxa II. Relatively large, flat platy tracheae, which are
expanded at their distal terminations, originate in the anterior region
of the genital vestibule and end above coxae II in Trachymolgus
(Grandjean, 1938).

Immature stages can be distinguished by the number of genital
setae and genital di-^cs. Protonymphs have one pair of genital
discs, deutonymphs have t^vo pairs, and tritonymphs and adults
have three pairs. Genital setae also increase in number in the
progressive instars; protonymphs have one pair (Biscirus silvaticus
has none), deutonymphs have two pairs (Cijfa latirosfris has one
pair), tritonymphs have four to eight pairs, and the adults possess
more genital setae than their respective tritonymphs.

A Revision of the Mite Family Bdellidae 361

SYSTEMATIC RELATIONSHIPS

Except for occasional oversights, there has been little question
among acarologists as to what constitutes the group commonly
known as the "snout mites." These mites have been identified as
being soft-bodied, predaceous mites with an elongated, conelike
gnathosoma, chelate chelicerae, and long, modified palpi. In 1902,
Thor divided the Bdellidae, erecting a new family, the Cunaxidae,
for "snout mites" with palpi modified for grasping prey and re-
taining the name Bdellidae for the remainder of the group.

Until recently, bdellids could be easily differentiated from cunax-
ids by the presence of one or two long, tactile setae at the apices of
the palpi. Although this character has been shown to occur in at
least one species of Cunaxidae (Atyeo, 1958), the families Bdellidae
and Cunaxidae still seem separable and the present paper deals only
with the former.

Bdellids are characterized as having well developed, chelate che-
licerae; padlike, rayed empodia; five-segmented, tactile palpi; two,
four, or five eyes; finely striated integument (except Trachymol-
gus); three pairs of genital discs; and usually more than one pair
of trichoboths. Cunaxids have chelicerae with the fixed digit
wanting; empodia reduced to a few rays; three to five-segmented
palpi usually modified for grasping prey and usually armed with
strong spines or apophyses; eyes present or wanting; integuments
usually with nonstriated plates; two pairs of genital discs; and
one pair of trichoboths.

In attempting to determine the probable relationships within
the Bdellidae, the genital tracheae and the ventral hypostomal
setae seem especially important. Two major groups can be rec-
ognized, one with well developed genital tracheae and two pairs
of ventral hypostomal setae, the, other without well developed
genital tracheae and with six to seven pairs of ventral hypostomals.
Within these two groups Grandjean (1938) placed genera showing
strong affinities into four subfamilies ( see key, p. 371 ) which appear
to be natural groups.

Except for Odontoscirus and the species of BdeJIa having inflated
cheliceral bases, the shape of the chelicerae, the position of the
cheliceral setae (except Neomolgus), and the unmodified chelae
are identical in both the groups lacking genital tracheae, the Odon-
toscirinae and the Bdellinae (text fig. la). In the subfamilies with
genital tracheae, the chelicerae are either elongated and have

362

The University Science Bulletin

Neomolgus

Bdellodes

Octobdellodes

Odontoscirus

Thnribdella

Bdella

More tnan two One or two pairs
pairs of clielic- of cheliceral se-
eral setae. tae.

Lateral propo-
dosomal setae
wanting.

Lateral propo-
dosomal setae
present.

Chelicerae nor-
mal, chelae un-
modified; lateral
lips small.

Chelicerae in-
flated, chelae
modified; lateral
lips large.

Posterior pseu-
dostigniata and
sensilla unmodi-
fied. I

Posterior pseu-
dostigmata and
sensilla modified.

Five pairs of
trichoboths; pal-
pal tibiotarsus
usually long,
cylindrical.

ODONTOSCIRINAE

Four pairs of
trichoboths (one
species with
three) ; palpal
tibiotarsus short,
truncated.

bdel

LINAE

fig. lb

Without well de-
veloped genital
tracheae; six to
seven pairs of
ventral hyposto-
mal setae.

With wcli devel-
oped genital tra-
cheae; two pairs
of ventral hypo-
stoinal setae.

Text fig. la. Probable relationsliips within the Bdellidae

A Revision of the Mite Family Bdellidae

363

Cyta

Trachymolgus

Integument armored. Integumentsoft, stri-
ated.

Spinibdella

Normal to inflated
chelicerae; one to
three pairs of tricho-
boths.

Biscirus
Two end setae.

Monotrichobdella
One end seta.

Palpal tibiotarsus
truncate; lateral pro-
podosomal setae
present.

Palpal tibiotarsus
cylindrical; lateral
propodosomal setae
wanting.

Elongate chelicerae;
four pairs of tricho-
boths.

CYTINAE

SPINIBDELIJXAE

Genital tracheae well
developed; two pairs
of ventral hyposto-
mal setae.

Text fig. lb. Probable relationships witliin the Bdellidae (Concluded) .

364 The University Science Bulletin

needlelike chelae (Spinibdellinae) or are greatly thickened with
massive chelae (Cytinae, except possibly Trachyniolgus), and the
anterior cheliceral seta is inserted on the distal fourth of the chelic-
era.

Only a few characters are available for generic separation (text
figs, la, lb). Although genera separated on these characters appear
to form natural groups, one diflFerence does occur that cannot be
evaluated. In the Odontoscirinae, only three species have lateral
propodosomal setae; Octobdellodes gen. nov. has been erected to
include two of these species. The third undoubtedly belongs to
ThoribdeUa, as it has highly modified posterior sensilla. Except
for this single species of ThoribdeUa, the presence or absence of the
lateral propodosomal setae is a stable generic character, therefore,
the exact significance of one anomalous species cannot be ascer-
tained.

Internal apodemes on the dorsal propodosoma occur in species
of the Bdellinae and the Cylinae. The large rectangular to reni-
form-shaped thickening of the integument varies according to the
species in all groups, but it is best developed in some members of
the Odontoscirinae. Cuticular deposition can only be considered
to be a continuum, the amount seen in any species depends not
only on the individual, but also on the optical equipment, the
method of clearing the specimen, and the nature of the mounting
medium. Hoploscirtis Thor, 1937 and Hoplomolgus Berlese, 1923
were erected for species of Bdellodes ( = Scirus ) and Neomolgus
( rr Molgus) in which the reniform thickenings (dorsal shields of au-
thors) could be readily detected. The distinguishing character
for both genera is apparently based on an extreme modification
of a continuous character, cuticular deposition.

An increased number of cheliceral setae appears only in Neo-
molgus and previously, Bdellodes has been considered to be the
only genus having one cheliceral seta. However, this character is
not sufficient to distinguish Bdellodes, as members of this genus
have been discovered with two cheliceral setae, and in undescribed
species of ThoribdeUa from Australia, the posterior cheliceral seta
is vestigial in one species and wanting in another species.

A similar modification of the lateral lips of the hypostome occurs
in both major groups of bdellids. In Cijta and Odontoscirus, these
structures are prominent and heavily fringed apically. In other
genera, the lateral lips are small and the cuticular fringe is minute
(fig. 2).

A Revision of the Mite Family Bdellidae 365

INTRASPECIFIC VARIATION

A preliminary investigation of intraspecific variation demonstrated
a constancy of characters, especially within local populations. Al-
though evidence for subspecies exists in some species, until more
extensive material has been gathered, it is judged advisable not to
name them.

The variation found in species of wide distribution is striking.
There appears to be a positive correlation between increase in body
size and decrease in temperature. Generally the larger species or
individuals of a given species of Cyfa, Bdella and NeomoJo^us occur
in the colder regions. Concrete data on this apparent geographical
variation is presented under a few of the widespread species for
which series from widely separated localities are available.

A few characters, such as striation patterns and gnathosomal struc-
tures may be unique for a species, or may be similar in more than one
species. The definitive character for species differentiation appears
to be the number, type and relative positions of the sensory setae
of the legs. Except for the attenuate sensory setae located on the
dorsal surfaces of the genua and proximal halves of the tibiae, spe-
cialized setae are extremely stable. Although these setae tend to be
in slightly different positions in each specimen, their relative posi-
tions are the same. To illustrate this phenomenon, the positions of
the sensory setae on tarsus I were plotted for thirty-eight specimens
of Bdella longicornis. In figure 71, the stippled area surrounding
the bases of each seta represents a composite of all the positions in
which the seta occurred. Generally, all the setae migrated either up
or down the tarsus, thus, even though the exact positions had
changed, each seta was approximately in the same relative position
to the other sensory setae.

TAXONOMY

Historical Account

Excellent historical accounts have been presented by Thor ( 1931 ),
Oudemans (1937), and Vitztlium (1931a). This discussion will
therefore be limited to the important systematic questions that have
been created in the past.

Latreille (1795) originally described the type genus Bdella as
having "Deux antennules filiformes, longues, coudces, de quatre ar-
ticles, dont le premier et le dernier fort longs: celui-ci termine par
deux poils. Trois valvules reunies, formant un bee advance, alonge
et conique . . ." "La pince rouge" Geoffroy, 1762, was selected

366 The University Science Bulletin

by Latreille for the type species and the description was based pri-
marily on Geoffroy's ilkistration. In the following year, Latreille
(1796) synonymized "La pince rouge" Geoffroy, 1762, with Acarus
longicornis Linnaeus, 1758. Controversy arises in the interpretation
of the relative lengths of the two longer palpal segments, the femur
and tibiotarsus. Latreille's description is ambiguous on this point,
as he states only that the two segments are strongly elongated.

Geoffroy's (1762) figure depicts "La pince rouge" as having a
four-segmented palpus: trochanter, femur (basifemur and telo-
femur), genu, and tibiotarsus. The palpal apices are shown to be
expanded distally, obliquely truncate, with two subequal end setae
slightly longer than the femora, and the tibiotarsi approximately two
thirds the length of the femora. "La pince rouge" clearly belongs
to the genus Bdella, sensu Thor (1931a). When Latreille's descrip-
tion is compared with Geoffroy's figure, the description loses its
ambiguity and the following synonymy becomes probable: Acarus
longicornis L., 1758 = "La pince rouge" Geoffroy, 1762 = Bdella
longicornis Latr., 1795 = Bdella longicornis, sensu Thor, 1931a.

Vitzthum ( 1931 ) interprets Latreille's description as meaning that
the longer segments, the femur and the tibiotarsus, are approxi-
mately equal in length, thereby placing Bdella in the Odontoscirinae,
sensu Thor (1931a) and Grandjean (1938). To the present author,
Vitzthum's interpretation seems incorrect.

Hermann ( 1804 ) erected the genus Scirus for ". . . les Mites
dont Linne et Geoffroy n'ont connu qu'une seule espece, appalee
par le premier acarus longicornis. . . ." His type species, Scirus
vulgaris, is figured as having short, widened palpal tibiotarsi and is
considered to be synonymous with Acarus longicornis L. (Thor,
1931a; Oudemans, 1937; non Vitzthum, 1931). The genus Scirus,
sensu Hermann must therefore be synonymized with Bdella, sensu
Thor.

A second species described by Hermann (1804), Scirus longiros-
tris, is figured as having the terminal palpal segment cylindrical,
approximately as long as the femur and the end hairs approximately
the length of the tibiotarsus, a condition commonly found only in
the Odonotscirinae. S. longirostris has been regarded as the tyj^De
species for Scirus, sensu Thor (non Hermann, 1804), but as Scirus
and Bdella are isogenotypic, the name Scirus is invalid. Oudemans
(1937) recognized this inconsistency and proposed the name
Bdellodes to replace Scirus, sensu Thor.

A Revision of the Mite Family Bdellidae 367

Characters and Descriptive Methods
To simplify the species descriptions, characters are presented in
an abbreviated form. Important morphological features used in the
descriptions are discussed fully in the morphology section. For
species being redescribed, the intraspecific variations in measure-
ments and counts ( e. g., setae ) observed in the study specimens are
expressed as ranges immediately following the character being de-
scribed. For new species, variations are noted in the remarks sec-
tion following each description.

A phase contrast microscope was used throughout this study. A
grid in the eyepiece was used to draw the figures to scale.

Following the same sequence as will be found in the formal
descriptions that follow, the characters and descriptive methods are
explained in the following section.

Color in life, which has been extensively used as a taxonomic char-
acter by earlier workers, is apparently dependent on body contents
in most species. Although no experiments were undertaken to
demonstrate that bdellids change color on different diets, the author
noted that Cijta coenilipes, the only species with a purple integu-
ment, in life varied from purple to brown with yellow blotches.
A few species have black, subcutaneous spots beneath the lateral
eyes, but these disappear after the specimens have been treated
with lacto-phenol clearing solution.

Length, including gnathosoma is a highly variable measurement.
Nongravid females and males are approximately the same size, but
the hysterosoma of gravid females becomes greatly distended,
thereby creating an erroneous impression of the species size.

Palpus. Illustrations are of the median aspect of the left palpus.
Measurements of the palpal segments were taken from the dorsal
surfaces and are given in the following order: trochanter (I), basi-
femur (II), telofemur (III), genu (IV), tibiotarsus (V), dorsal end
seta (des), and ventral end seta (ves). Variations in the number of
setae are indicated in the remarks section following each description.

Chelicera. The shape of the chelicera may be normal, which
means that it is approximately three to five times as wide at the
base as at the thinnest portion ( fig. 12 ) . Inflated chelicera indicates
that the width at the base is more than seven times greater than at
the thinnest portion (fig. 13), and elongated chelicera has sides that
are approximately parallel (figs. 32-35). The chelae in the majority
of the genera are small and the ventral, movable difrif ciVl^V.^hnrx^d
(fig. 7h). Smooth chelae means that the blades lack teeth on the

368 The University Science Bulletin

inner surfaces. In the Odontoscirinae, the inner surface of the
movable digit may be flattened subapically. This flattened surface
extends proximally to the level of the tip of the fixed digit (fig. 8).
The length of a chelicera is measured from the base to the apex of
the longest digit.

Hyposiome. As viewed from the ventral aspect (fig. 2), the
gnathosoma consists of a rectangular base carrying the palpal
articulations at the anterolateral angles, and an anteriorly projecting
hypostome. The latter structure consists of a long, tapering buccal
cone, and two small, distally fringed lateral lips. The gnathosomal
base is always transversely striated, but the buccal cone may be
striated or nonstriated. In the latter case, the integument may
appear to be granulated. In adults, there are two, six or seven pairs
of large setae in two longitudinal series on the ventrolateral surfaces
of the hypostome that extend from the anterior margin of the
gnathosomal base to the proximity of the lateral lips. These setae,
the ventral hypostomals, are indicated as the vh series and are
numbered consecutively from the base, thus, the proximal pair of
setae is vh 1. The positions of the ventral hypostomal setae may be
important in future taxonomy, therefore, relative positions are in-
dicated in the descriptions. Two dorsal hypostomal setae may be
present beneath the cheliceral bases. If present, these setae are
inserted approximately above the palpal articulations. The lengths
of the dorsal h\'postomal setae are compared to the lengths of the
setae comprising the ventral series.

Propodosoma. The integumental striae of this area, although not
always specific, appear in different structural types and patterns.
The breaks along the crests of these integumental ridges vary in
frequency and regularity and on this basis have been divided into
three categories. Sparsely broken striae (fig. 136) indicates that
the breaks occur at long and irregular intervals. Coarsely broken
striae (fig. 151) divide at regular intervals with the lengths of the
uninterrupted crest approximately six to eight times longer than the
breaks. Finely broken striae ( fig. 133 ) indicates that the undivided
lengths of the crests are only two to three times longer than the
breaks. Although it might be expected that many intergrades would
occur between these arbitrary groups, this is not the case. Inter-
grades occur in only one or two instances, and these are between
the coarsely and finely broken striation groups.

The lateral propodosomal setae (figs. 1 d, 133 c) are inserted
lateral to a line connecting the dorsal sensilla on each side. The

A Revision of the Mite Family Bdellidae 369

presence or absence of this pair is considered to be of generic
significance (except Thoribdella). The median propodosomals
(fig. 1 f, 133 /) are ahvays present and are inserted in the interval
between the posterior sensilla.

On each side of the body, the eyes may be approximate or distant
and the distance between the eyes is compared to the diameter of
either the anterior or posterior eyes. In Bdella and Spinibdella, the
direction of the striae in the interocular interval may be longi-
tudinal, thus connecting the eyes, or transverse.

Dorsal hysterosoma. The interval between an internal humeral
and an internal dorsal seta (fig. 1 h, /) is considered to be the first
interspace and is used as a comparison with the length of the
humeral seta.

Anal region. The border striae are those striae which parallel the
entire length of the anal cleft or those which bend sharply latcrad
near the center of the cleft. Setae inserted in the region of the
parallel striations are the anal setae (fig. 2 /'). A pair of postanal
setae (fig. 1 o) flanks the dorsal termination of the cleft. Other
setae surrounding the anal cleft are considered to be the paranal
setae.

Genital region. Two membraneous plates or flaps cover the
genital vestibule, each of which has a regular or irregular row of
genital setae (figs. 2 i, 178 a). Setae surrounding these plates are
the paragenital .setae (figs. 2 g, 178 b), and although a few of these
setae occur between the coxae, no satisfactory division could be
made to separate these setae into groups of paragenitals and inter-
coxals. In the Cytinae, an unpaired median seta occurs immedi-
ately anterior to the genital flaps (fig. 178 c).

Setae inserted on the ovipositor are indicated by position. Those
surrounding the apex are termed the subapical setae, and those
slightly distal to the midlength of the ovipositor are termed the
postmedial setae.

The large, heavily sclerotized amphioid sclerites of the male
each bears a peripheral row of setae of various lengths and shapes
(figs. 169-177). In Spinibdella and Cyta, the differences in the
peripheral setae are believed to be specific. In other genera, setal
differences could not be evaluated. However, in this latter group
of genera, the setae are divided into distinct groups and are re-
corded as such in the descriptions. For example, "amphioid scle-
rites with 4, 1, 3, 2" would indicate that in the peripheral row of

370 The University Science Bulletin

setae, a group of four setae are anterior, then a single seta, then a
group of three setae, and finally near the caudal termination of
the sclerite, two setae.

Legs. Small or minute lateral rays on the claws refers to a row
of tiny spines, each of which is approximately the width of the
claw. Large lateral rays are similar in structure to the above, but
are less numerous and more than three times the width of the
claw (fig. 70). Measurements of the legs are given in microns
for the tibia and tarsus (excluding the pretarsus) of legs I and II.
The chaetotaxy is given for each segment, starting with coxa I.
If only one type of seta occurs on more than one segment of a
series (e. g., coxae), this is indicated in an abbreviated form. For
example, "coxae I-IV, 5, 4, 6, 4 tactile setae" means that only tactile
setae occur on the coxae and coxa I has five tactile setae, coxa II
has four tactile setae, etc. When segments have more than one type
of seta, each segment is given individually. The different types
of setae found on the legs are illustrated in figs. 179-197.

Description of the Family Bdellidae
Bdellei Duges, 1834, Ann. Sci. Nat., ser. 2, vol. 1, Zool., p. 21.

Medium to large mites (approximately 0.5 to 3.5 mm.) with
finely striated integument (Trachymolgus armored); subcutaneous
shields may be present on the propodosoma; conelike gnathosoma
formed by elongated, chelate chelicerae bearing one or more setae,
and a ventral hypostome bearing two, six, or seven pairs of con-
spicuous ventrolateral setae and two pairs of minute setae inserted
on the lateral lips; geniculate palpi five-segmented, trochanter and
genu short, femur divided distally into a short telofemur and a long
basifemur; tibiotarsus of various lengths and bears apically two long
setae (one in Monotrichobdella) ; tracheal openings near cheliceral
bases; idiosoma divided dorsally into propodosoma and hystero-
soma; four dorsal sensilla inserted in pseudostigmatic organs; two,
four, or five eyes present; legs eight-segmented (including short
pretarsus) with two claws and a padlike, rayed puvillus; maximally
five pairs of trichoboths; well developed genital tracheae may be
present; three pairs of genital suckers; anal cleft terminal; sexes
similar. The subfamilies erected by Grandjean (1938) are incor-
porated in the following generic key.

A Revision of the Mite Family Bdellidae 371

Key to the Genera of the Family Bdellidae

1. \'enter of hypostome with six or seven pairs of strong setae and
two pairs of small adomal setae; without well developed genital

trach

eae

Venter of hypostome with two pairs of strong setae and two pairs
of small adomal setae; with well developed genital tracheae 7

2. Trichoboth absent on tibia II ( Bdellinae ) Bdella p. 372

Trichoboth present on tibia 11 ( Odontoscirinae ) 3

3. Each chelicera vwth more than two setae, usually eight to twenty

Neomolgus p. 389
Each chehcera with one or two setae 4

4. Posterior pseudostigmata goblet-shaped; posterior sensilla reduced
in size, usually much shorter than median propodosomal setae

Thoribdella p. 394
Posterior pseudostigmata simple; posterior sensilla not reduced in
size and longer than median propodosomal setae 5

5. Lateral propodosomal setae present OctobdeUodes p. 407

Lateral propodosomal setae absent 6

6. Chela with inner surface of movable digit straight and bearing
one or more small teeth; cheliceral bases inflated; lateral lips

of hypostome prominent Odontoscirus p. 386

Chela with movable digit sickle-shaped and bearing one or no
teeth; cheliceral bases not inflated; lateral lips of hypostome
as in other genera Bdellodes p. 412

7. Cheliceral bases normal to inflated, chela with movable digit
sickle-shaped; unpaired median seta immediately anterior of

genital opening ( Cytinae ) 8

Chelicerae elongated; chelae with digits reduced, needlelike;
unpaired median seta, if present, between coxae IV

( Spinibdellinae) 9

8. Integument soft, striated; chelae massive; unpaired median eye

between anterior sensilla Cyta p. 416

Integument armored, pitted; chelae not massive; without unpaired
median eye Trachymolgus p. 423

9. Lateral propodosomal setae present; palpal tibiotarsus expanded

distally ( truncated ) SpinihdcUa p. 424

Lateral propodosomal setae absent; -palpal tibiotarsus cylindrical,
elongated 10

10. Palpal tibiotarsus with two long apiacl setae Biscinis p. 435

Palpal tibiotarsus witli one long apical seta ... Monotrichobdella p. 438

Bdellinae Grandjean, 1938

The Bdellinae, containing only the genus Bdella, is characterized
by having six pairs of ventral hypostomal setae, four pairs of tricho-
boths {Bdella mexicana has three pairs), and undeveloped genital
tracheae. No exceptions were found in the material examined;
however, Vitzthum ( 1943 ) reports an undescribed species of
Bdella which has one pair of weakly developed genital pouches.

372 The University Science Bulletin

Genus Bdella Latreille

Bdella Latreille, 1795, Magasin encyclopcdique, on Journal des Lettres et des
Arts, Paris, vol. 4, p. 18. (Type: Acarus longicarnis Linnaeus, 1758, by
subsequent identification, Latreille, 1796, Precis des Caracteres goneriques
des Insectes, disposes dans un Ordre naturel, par [Pierre Andre] Latreille,
Paris, p. 180).

CJieUfer Geoffroy, 1762, llistoire abregee des Insectes, qui se trou\ent aux
En\irons de Paris; dans lacjuelle ces Animaux sont ranges suivant un Ordre
mrtiiodique, par Etienne Louis Geoffroy, vol. 2, p. 617. (Type: Chelifer
totus ruber, antennis extremo bisetis: Pince rouge Geoffroy, 1762 [= Acarus
longicornis L,.] [monobasic].)

Scirus (Ciron) Hemiann, 1804, Memoire apterologique. Ouvrage couronne en
1790 par la Societe d'Histoire naturelle de Paris, Public par Frederic-Louis
Hammer, Strasbourg, p. 60. (Type: Scirus vulgaris Hermann, 1804
[^Acarus longicornis L.] [first included species].)

Bdellidium Oudemans, 1929, Ent. Ber. Nederl. Ver., vol. 7, p. 449 (new
synonym). (Type: Scirus vulgaris Hermann, 1804 [^Acarus longicornis
L.] [by original designation].)

Caenobdella Oudemans, 1937, Kritisch Historisch Overzicht der Acarologie,
Leiden, vol. 3, part C, p. 1227 (new synonym). (Type: Bdella crassipes
C. L. Koch, 1839 [by original designation].)

In addition to the subfamily characters, the palpal tibiotarsus is
truncate and considerably shorter than the palpal basifemur; the
subequal end setae are as long as or longer than the palpal femur.
Normal to inflated chelicerae bear two setae inserted on the proxi-
mal three fourths of their lengths, and the small chelae have sickle-
shaped movable digits, each of which may have one small tooth.
The dorsal propodosoma with four pairs of eyes lateral to the un-
modified posterior pseudostigmatic organs. The podocephalic canal
is an external groove (Grandjean, 1938).

The genus Caenobdella Oudemans is based on the inadequate
description of Bdella crassipes Koch, 1839. Koch's illustration shows
five trichoboths on the right side of the animal and four trichoboths
on the left side. The trichoboths of legs III, one pair inserted on
tibiae III, one on the right tarsus III, are the distinguishing features
of Caenobdella. In this author's opinion, these discrepancies repre-
sent mistakes made by Koch, and therefore, should not be con-
sidered to represent significant structures.

Key to the Species of Bdella

1. Distance between anterior sensilla bases less than between pos-
terior sensilla; striae of propodosomal shield, if longitudinal, not

directed between anterior sensilla 2

Distance between anterior sensilla bases greater than between pos-
terior sensilla; striae of propodosomal shield longitudinal, directed
between anterior and posterior sensilla 5

2. Propodosomal shield with all striae convex caudally 3

Propodosomal shield with anterior striae convex caudally, posterior
striae longitudinal, directed between posterior sensilla 4

A Revision of the Mite Family Bdellidae 373

3. Shield with finely broken striae; palpus with thirteen or more setae
on basifemur, seven setae ( including end setae ) on tibiotarsus

longicornis p. 373
Shield with sparsely broken striae; palpus with eight to eleven
setae on basifemur, six setae (including end setae) on tibiotarsus

muscorum p. 375

4. Telofemur, genu, and tibia of leg I approximately equal in length,

genu I without duplex seta tropica p. 378

Tibia I two times longer than genu I or telofemur I; genu I with

duplex seta longistriata p. 380

5. Tarsus IV with trichoboth; dorsal hysterosomal setae branched

distally; transverse striae between eyes distincta p. 381

Tarsus IV without trichoboth; dorsal hysterosomal setae nude or
plumose; longitudinal striae between eyes mexicana p. 383

Bdella longicornis Linnaeus

(Figs. 7, 12, 37, 69-72, 133-135, 169)

Acarus longicornis Linnaeus 1758, Systema Naturae, 10th ed., p. 618.
"Chelifer totus ruber, antcnnis extreme bisetis; Pince rouge" GeoflFroy, 1762,

Hist. Abr. Ins., vol. 2, pp. 618-619.
Scirus vulgaris Hermann, 1804, Mein, Apt., p. 61.
Bdella anguinesetosa Ewing, 1910, Univ. Stud., Univ. Illinois, vol. 3, no. 6, p.

72 (new synonym).
Bdella tessellata Ewing, 1913, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 112

(new synonym).

This widespread species is closely related to Bdella muscorum
Ewing, 1909, and to Bdella iconica Berlese, 1923. B. longicornis
can be distinguished from B. muscorum by having seven setae (in-
cluding two apical setae) on the palpal tibiotarsus rather than six,
and in having the interlacing secondary apodemes of the dorsal
propodosomal shield lateral to the bell-shaped primary apodemes
(fig. 133) rather than mesad (fig. 136). The palpal tibiotarsus of
B. iconica is identical to that of longicornis, but the striae of the
propodosomal shield are convex caudad in longicornis rather than
being directed between the median propodosomal setae.

Female: Color in life, deep pink to red, with irregular brown
to dark blue blotches. Body ovoid, weakly constricted; length,
including gnathosoma, 1160 [j. (850-1475 ix). Gnathosoma: Length,
320 [JL (209-396 \i.); palpus (fig. 37) with tibiotarsus extending be-
yond hypostome; measurements: I, 15 a (11-16 [i); II plus III,
327 [X (142-371 ix); IV. 30 [x (20-38 [;.); V, 95 '^ (54-115 [j.); des,
243 jx (187-464 [x); ves, 195 [x ( 141-367 (x). Chelicera ( figs. 7, 12 )
normal, finely striated, 300 [x (193-423 jx) in length; movable digit
smooth, equal in length to fixed digit; setae as in fig. 12. Hypo-
stome striated; vh 1 and vh 2 form transverse row between palpal
articulations; dorsal hypostomals wanting. Dorsal propodosoma

374 The University Science Bulletin

(figs. 133-135): Striae finely to coarsely broken; lateral propodo-
somals nude, 57 [x (43-100 \).); median propodosomals nude, 67 \).
(57-128 \>.) in length; eyes separated by diameter of anterior pair;
distance between anterior sensilla, 74 [>. (56-143 [j. ); large pore
anterolaterad to lateral propodosomal seta. Dorsal hijsterosoma:
Setae nude; internal humeral, 70 [j, (57-116 [j.) in length, approxi-
mately the length of first interspace; external humeral, 99 \x (70-
184 [jl) in length; sacrals and clunals in gently curving rows. Anal
region: Border striae parallel; one pair of anals; 5 pairs of paranals;
postanals anterior to cleft, longer than external clunals. Genital
region: Each genital plate with 7 (6-8) equal genital setae in ir-
regular linear arrangement; 9 (10) pairs of paragenitals, anterior
pair not between coxae IV; genital discs small, equidistant, in
posterior two thirds of vestibule; ovipositor with 10 subapical and
10 postmedial setae. Legs (figs. 69-72): Claws with 4-5 lateral
rays, 1 row of minute rays each; measurements: tibia I, 114 a (65-
180 \i.); tarsus I, 146 [;. (100-201 [x); tibia II, 87 [x (52-143 ij.); tar-
sus II, 125 \>. (83-180 ij.). Chaetotaxy: coxae I-IV, 6 (5), 6 (5),
5, 3 (4) tactile setae; trochanters I-IV, 1, 1, 2, 2 tactile setae;
basifemora I-IV, 13 (11-17), 11 (11-15), 11 (11-14), 6 (7) tactile
setae; telofemora I-IV, 11 (11-13), 11 (11-13), 11 (12), 9 (9-11)
tactile setae; genu I, 9 (8) tactile setae, 1 duplex seta, 3 (3-6) at-
tenuate sensory setae; genu II, 9 tactiles, 1 duplex, 1(2) attenuate
sensory setae; genu III, 9 tactiles, 1 duplex, 1 attenuate sensory
seta; genu IV, 10 (11) tactiles, 1 attenuate sensory seta; tibia I,
16 tactiles, 5 (5-9) attenuate sensory setae, 1 attenuate peg, tricho-
both; tibia II, 15 (13-15) tactile setae, 2 attenuate sensory setae,
one blunt sensory seta; tibia III, 16 ( 17 ) tactile setae, one atten-
uate sensory seta; tibia IV, 15 tactiles, trichoboth; tarsus I, 14 ( 16 )
ventrals, distal pairs plumose, 10 lateral and 2 dorsal tactile setae,
2 attenuate and 2 blunt sensory setae, 1 attenuate peg, dorsotermi-
nals minutely plumose; tarsus II, as in tarsus I, except lacking 2 at-
tenuate sensory setae and 1 lateral seta; tarsus III, 14 ventrals, distal
pairs plumose, 9 laterals, trichoboth, dorsoterminals as in leg I;
tarsus IV, 14 ventrals, 7(8) laterals, trichoboth, 1 attenuate sensory
seta distal to trichoboth, dt 1 reduced to one seta, dt 2, 3 minutely
plumose.

Male: Identical to female; amphioid sclerites (fig. 169) with 7, 1,
2 setae each.

Type: Europe.

Location of type: Unknown.

A Revision of the Mite Family Bdellidae 375

Remarks: Two of H. E. Ewing's types available for study from
the U.S. National Museum were: Bdella anguinisetosa (trito-
nymph), June 5, 1905, H. E. Ewing; and Bdella tessellata (female),
Portage, Wisconsin, September 2, 1909, H. E. Ewing, under an old
piece of bark.

In addition, approximately 250 specimens were examined, which
included material from: Barro Colorado Island, Panama Canal
Zone, Costa Rica, Cuba, Mexico (Oaxaca, Distrito Federal, Mex-
ico, San Luis Potosi), United States (California, Texas, Utah, Ar-
kansas, Kansas, Illinois, Tennessee, Florida, Missouri, Michigan,
New Hampshire, Vermont), and Nova Scotia.

The numbers of setae on the palpal segments are: basifemur, 14
( 13-15); telofemur, 1; genu, 4; tibiotarsus, 4 tactile setae, 1 attenuate
sensory seta, and two long apical setae. The length of the solid
tactile seta on the dorsal surface of the palpal tibiotarsus is slightly
longer, to more than twice the length of the attenuate sensory seta.

The variation in total length is considerable, but more striking is
the variation in cuticular deposition shown by the internal apodemes
of the dorsal propodosoma (figs. 133-135). Geographical variation
could not be demonstrated in this species. Within one area, indi-
vidual mites could be found which exhibited all combinations of
the variable characters.

The illustrations were prepared from females from: West slope

of Cortez Pass, Mt. Popocatepetl, Mexico, Mexico, August 11, 1954,

W. T. Atyeo, moss sample off tree (Berlese funnel extraction), alt.

11,500 ft.

Bdella musconim Ewing

(Figs. 39, 73, 74, 136-139)

Bdella muscorum Ewing, 1909, Canadian Ent., vol. 41, no. 4, pp. 124-125.
Bdella lata Ewing, 1910 {non Koch & Berendt, 1854, fossil Bdella?), Univ.

Stud., Univ. Illinois, vol. 3, no. 6, p. 69.
Bdella subnigra Ewing, 1910, Univ. Stud., Univ. Illinois, vol. 3, no. 6, p. 73

(new synonym).
Bdella muscorum var. minnesotcnsis Ewing, 1913, Bull. Anier. Mus. Nat. Hist..

vol. 32, p. 113 (new synonym).
Bdella recens Ewing, 1937 {pro, Bdella lata Ewing, nom praeocc.), Univ.

Toronto Stud., Geological Ser., no. 40, p. 57 (new synonym)-

This species is closely related to Bdella longicornis, and due to
many superficial similarities, these two species may be easily con-
fused. In addition to differences in the chaetotaxy of the legs, B.
muscorum can be easily differentiated by the palpal chaetotaxy, the
basifemur having eight to eleven setae and the tibiotarsus having a
total of six setae. The palpus of B. longicornis has thirteen or more
setae on the basifemur and seven setae on the tibiotarsus.

376 The University Science Bulletin

Female: Color unknown. Body ovoid, weakly constricted; length,
including gnathosoma, 825 [i (710-1170 ;;.) Gnathosoma: Length,
224 [x (206-331 \l); palpus (fig. 39) with tibiotarsus extending
beyond hypostome; measurements: I, 15 [t. (13-18 [x); II plus III,
135 [X (133-198 ^); IV, 26 ix (20-35 [j.); V, 60 [x (55-99 [x); des, 175 [x
(110-220 11.) ; ves, 129 ii (129-182 [x). Chelicera normal, finely
striated, 215 [x (207-300 [x) in lengch; chela smooth, fixed digit blunt,
slightly shorter than movable digit; distal seta extending to base of
chela, proximal seta extending to insertion of distal seta; setae in-
serted as in fig. 12. Gnathosoma striated; vh 1-2 in strongly curved
transverse row between palpal articulations, vh 2, 3, 5, 6 progres-
sively more widely separated and form 2 longitudinal series, vh 1, 4
laterad of series; dorsal hypostomal setae wanting. Dorsal pro-
podosoma (figs. 136-139) : Striae sparsely broken; lateral propodoso-
mals nude, 57 [j. (47-78 (x) in length; median propodosomals nude,
74 [X (71-99 [x) in length; eyes separated by distance equal to
diameter of anterior pair, interval between eyes with longitudinal
striae; distance between anterior sensilla, 70 [x (61-97 jx). Dorsal
hysterosoma: Setae nude; length of internal humeral, 79 [x (79-98 [x),
approximately three fourths of first interspace; external humeral,
105 [;. (92-124 [x) in length; sacrals and clunals in gently curvdng
transverse rows. Anal region: Border striae parallel, 2 pairs of anal
setae, 5(4) pairs of paranals anterior to termination of cleft, longer
than external clunals. Genital region: Each genital plate with 8(9)
small equal genital setae in linear arrangement; 10 pairs of para-
genitals, anterior pair not between coxae IV; genital discs small,
approximately equidistant, in posterior three fourths of vestibule;
ovipositor with 10 subapical and 8 postmedial setae. Legs ( figs. 73,
74 ) : Claws with 4-6 lateral rays, one row of minute rays each; meas-
urements: tibia I, 74 [x (64-100 ix); tarsus I, 90 -j, (85-124 [x); tibia II,
53 [X (45-82 [x); tarsus II, 75 [x (71-108 [x). Chaetotaxy: coxae I-IV,
5(6),6(5),5(7),4(3) tactile .setae; trochanters I-IV, 1, 1 (2),
2. 2 tactile setae; basifemora I-IV, 10 (8-14). 8 (7-10), 9 (8), 5 (6)
tactile setae; telofemora I-IV, 7 (8-11), 7 (6-10), 7 (8-9), 8 (7)
tactile setae; genu I, 6 (6-8) tactile setae, 1(2) attenuate sensory
seta(e), 1 duplex seta; genu II, 6 (6-8) tactiles, 1 duplex; genu III,
6 (6-8) tactiles, 1 duplex; genu IV, 8 tactiles, 1 attenuate sensory
seta; tibia I, 14 (13) tactiles, 4 attenuate sensory setae, 1 peg,
trichoboth; tibia II, 11 tactiles, 2(1) attenuate sensory .seta(e), 1
blunt sensory seta; tibia III, 12 (10-14) tactiles, 1 attenuate sensory
seta; tibia IV, 13 (11-12) tactile setae, trichoboth; tarsus I, 13 (14)

A Revision of the Mite Family Bdellidae 377

plumose ventrals arranged in 2 rows, 8 lateral and 2 dorsal tactiles,
2 attenuate and 2 blunt sensory setae, 1 peg, dt 1 hollow, nude, dt 2,
3, solid, plumose; tarsus II, 10 (11-12) plumose ventrals, 6 lateral
and 2 dorsal tactiles, 2 blunt sensory setae, 1 peg, dorsoterminals as
in tarsus I; tarsus III, 12 (11) ventrals, 7 (6-8) laterals, trichoboth,
dt 1 solid, nude, dt 2, 3, solid, plumose; tarsus IV, 12 ( 11 ) ventrals,
6 laterals, 1 attenuate sensory seta, trichoboth, dt 1 reduced to one,
solid, nude seta, dt 2, 3, solid, plumose.

Male: Identical to female; amphioid sclerites with 6, 1, 2 setae
each.

Type: Female, Muncie, Illinois, June 16, 1908, H. E. Ewing, in
moss.

Location of type: The United States National Museum.

Remarks: Three additional type specimens were examined and
found to be conspecific with B. muscorum; these were: Bdella
recens Ewing, 1937 {pro, B. lata, Ewing, nom praeocc.), sex un-
known (incomplete specimen), Mahomet, Illinois, April 17, 1908,
H. E. Ewing; B. .stdjnigra Ewing, 1910, female, Mahomet, Illinois,
April 17, 1908, H. E. Ewing, in moss; B. muscorum var. minnesoten-
sis Ewing, 1913, female, Minnesota, April 24, 1900. Additional
material studied included specimens from: The United States (Cali-
fornia, Colorado, New Mexico, Kansas, Arkansas, Tennessee, Michi-
gan, Illinois, Maryland), Alaska (Point Barrow, Chandler Lake
Region, Umiat, District of Mackenzie), Germany (Ost-Holstein),
Czechoslovakia, and Iceland.

In the Old World specimens, the mesal margins of the dorsal
propodosomal apodemes are more highly developed than the lateral
margins, whereas in the New World material, the reverse is true
(fig. 136). However, in either condition, there is little variation in
the characteristic shape of the apodemes.

As indicated in the description, the chaetotaxy of the legs is rela-
tively constant. The chaetotaxy of the palpus is as follows: basi-
femur, eight to eleven tactile setae, usually ten; telofemur, one seta;
genu, four setae; tibiotarsus, three tactile setae, one attenuate sen-
sory seta, and two long apical setae. The dorsal sensory seta and
the dorsal tactile seta fluctuate considerably in actual and relative
lengths. The tactile seta may be slightly longer, or as in the Ice-
landic specimens, almost twice the length of the sensory seta. Draw-
ings of a female from Douglas Lake, Cheboygan Co., Michigan, June
24, 1957, R. E. Beer, moss (Berlese funnel extraction).

378 The University Science Bulletin

Bdella tropica, sp. nov.
(Figs. 40,77, 78, 140)

This species is closely related to Bdella distincta, but has long,
nude, dorsal hysterosomal setae rather than short, distally branched
setae, and has a different propodosomal striation pattern (figs. 140,
141). Although this new species does not appear to be closely
related to Bdella grandjeani Thor (1931b) from Tanganyika, the
propodosomal striae form similar patterns in both species.

Female: Color in life unknown. Body narrowly ovoid, weakly
constricted; length, including gnathosoma, 810 \i.. Gnathosoma:
Length, 211 \i; palpus (fig. 40) short, only apex of tibiotarsus ex-
tending beyond hypostome; measurements: I, 12 [i; II plus III, 121
[x; IV, 24 pi,; V, 45 [Ji; des, 175 [x; ves, 153 [i.. Chelicera normal, finely
striated, 204 [jl in length; chela smooth, fixed digit attenuate, ex-
tending slightly beyond movable digit; distal seta extending to base
of chela, slightly longer than proximal seta; setae inserted as in
Bdella longicornis (fig. 12). Gnathosoma faintly striated; ventral
setae in two longitudinal series, vh 1 between palpal articulations,
distance between vh 1 and vh 2 equal to distance between vh 5 and
vh 6, distance between vh 2 and vh 3 equal to one half distance be-
tween vh 3 and vh 4 or vh 4 and vh 5; dorsal hypostomal setae deli-
cate, same length as proximal pair of ventral setae. Dorsal propo-
dosoma ( fig. 140 ) : Striae finely broken; lateral propodosomals nude,
56 [JL in length; median propodosomals nude, 125 \t. in length; eyes
approximate, separated by 3 transverse striae; distance between an-
terior sensilla, 92 jj.. Dorsal hysterosotna: Setae nude; length of in-
ternal humeral, 75 [x, approximately five sixths of first interspace;
external humeral, 238 [x in length; sacrals and clunals in gently
curving transverse rows. Genital region: Each genital plate with
8 small, equal, genital setae in linear arrangement, 9 pairs of para-
genitals, anterior pair between coxae IV; genital discs small,
anterior pair near center of vestibule, posterior two pairs approxi-
mate, near caudal end; ovipositor with 10 subapical and 8 postme-
dial setae. Anal region: Border striae bending laterad; one pair
of anal setae anterior, one pair posterior to laterally-directed striae;
one pair of paranals; postanals flanking termination of cleft, shorter
than clunal setae. Legs (figs. 77, 78): Claws with 2-3 basal, lateral
rays and one row of minute rays each; measurements: tibia I, 51 [x;
tarsus I, 91 [x; tibia II. 38 \).; tarsus II, 79 ij.. Chaetotaxy: coxae I-IV,
6, 5. 6, 4 tactile setae; trochanters I-IV, 1, 1, 2, 1 tactile setae; basi-
femora I-IV, 8, 9, 7, 4 tactile setae; telofemora I-IV, 5 tactile setae

A Revision of the Mite Family Bdellidae 379

each; genu I, 5 tactile setae, 2 attenuate sensory setae, 1 duplex
seta; genua II-III, 6 tactile setae and 1 duplex seta each; genu IV,
5 tactile setae, 1 duplex seta; tibia I, 7 tactile setae, 3 attenuate
sensory setae, 1 attenuate peg, trichoboth; tibia II, 7 tactile setae,
2 attenuate sensory setae, 1 blunt sensory seta; tibia III, 8 tactile
setae, 1 attenuate sensory seta; tibia IV, 10 tactile setae, trichoboth;
tarsus I, 9 plumose ventral setae arranged in 2 rovi^s, 7 lateral and
2 dorsal tactile setae, 2 blunt and 2 attenuate sensory setae, 1 hol-
low, blunt peg, dt 1 hollow, pilose, dt 2, 3 solid, plumose; tarsus II,
8 plumose ventrals, 4-5 lateral and 2 dorsal tactile setae, 2 blunt
sensory setae, 1 peg, dorsoterminals as in tarsus I; tarsus III, 10
plumose ventrals, 6 laterals, trichoboth, dt 1 solid, nude, dt 2, 3
solid, plumose; tarsus IV, 10 plumose ventrals, 5 laterals, trichoboth,
1 attenuate sensory seta distal to trichoboth, dorsoterminals as in
tarsus III.

Male: Unknown.

Holotype: Female, Barro Colorado Island, Canal Zone, Panama,
July 27, 1956, Carl W. Rettenmeyer, bark moss (Berlese funnel
extraction ) .

Paratypes: Twenty-three females from Barro Colorado Island,
Canal Zone, Panama, collected by Carl W. Rettenmeyer by Berlese
funnel extraction with the following data: nine females, same data
as the holotype; eight females collected July 24, 1956, rotten stump;
three collected July 10, 1956, grass roots from clearing; two col-
lected August 1, 1956, soil and leaves; one collected March 26,
1956, rotten stump; one collected July 28, 1956, dirt and leaves.
Also, four females, Barro Colorado Island, May 12, 1956, C. W.
and M. E. Rettenmeyer, colony E:148, Eciton burchelli, refuse
deposit.

Location of types: The holotype and twenty-one paratypes are
deposited in the Snow Entomological Museum; two paratypes are
deposited at each of the following institutions: the United States
National Museum, the British Museum (Natural History), and the
South Australian Museum.

Remarks: Little variation is apparent in this species. The length,
including the gnathosoma, varies from 780 pi to 910 \i, and other
measurements vary proportionately, e. g,., the palpal femur, 112 tj. to
124 [JL. The number of tacitle setae on the leg segments vary from
one more, to one less, than the number indicated in the description
of the holotype. This species, collected only on Barro Colorado
Island, may be restricted to the tropical regions, hence the name
tropica. Drawings of the holotype.

380 The UNivERSiri' Science Bulletin

Bdella longistriaia, sp. nov.

(Figs. 38, 75, 76, 151, 170)

This species may be related to Bdella tropica, sp. nov., but can be
easily distinguished by the lack of long setae on the dorsal surface
of the palpal tibiotarsus.

Female: Color in life unknown. Body ovoid, weakly constricted;
length, including gnathosoma, 902 [jl. GnatJio.sonia: Length, 252 [x;
palpus (fig. 38) with tibiotarsus extending beyond hypostome;
measurements: I, 17 pi; II plus III, 190 \).; IV, 21 y.; V, 48 [x; des, 255
[x; ves, 204 \}.. Chelicera normal, finely striated, 274 [x in length; chela
smooth, fixed digit attenuate, slightly shorter than movable digit;
distal seta extending three fourths of distance to base of chela,
proximal seta extending one half the distance to distal seta; setae
inserted as in fig. 12. Gnathosoma striated; vh 1-6 in two longi-
tudinal series, vh 1 between palpal articulations, vh 1-5 approxi-
mately equidistant, vh 6 midway between vh 5 and apex; dorsal
hypostomal setae wanting. Dorsal propodnwina (fig. 151): Striae
sparsely broken; lateral propodosomals nude, 60 pi, in lensfth; median
propodosomals thickened, nude, 146 [j. in length; eyes separated by
distance equal to diameter of anterior pair, interval between eyes
transversely striated; distance between anterior sensilla, 92 \i. Dorsal
hysterosoma: Setae thickened, nude; length of internal humeral,
98 [JL, approximately one fourth longer than first interspace; external
humeral, 153 p. in length; sacrals and clunals in gently curving trans-
verse rows. Anal region: Border striae parallel; anal setae wanting;
2 pairs of paranals; postanals flanking termination of cleft, equal in
length to external clunals. Genital region: Each genital plate with
eleven equal setae in linear arrangement; 9 pairs of paragenitals,
anterior pair between coxae IV; genital discs small, widely sep-
arated, one pair at each end of vestibule, one pair near middle;
ovipositor with 10 subapical, 8 postmedial setae. Legs (figs. 75, 76) :
Claws with one row of small, lateral rays each; measurements: tibia
I, 81 [i; tarsus I, 108 [x; tibia II, 60 [x; tarsus II, 102 [x. Chaetotaxy:
coxae I-IV, 5, 4, 6, 5 tactile setae; trochanters I-IV, 1, 1, 2, 1 tactile
setae; basifemora I-IV, 11, 12, 9, 4 tactile setae; telofemora I-IV,
9, 9, 9, 6 tactile setae; genu I, 7 tactile setae, 1 duplex seta, 1 attenu-
ate sensory seta; genua II-IV, 8 tactile setae, 1 duplex seta each; tibia
I, 12 tactile setae, 2 attenuate sensory setae, 1 attenuate peg, tricho-
both; tibia II, 12 tactile setae, 1 attenuate and 1 blunt sensory seta;
tibia III, 9 tactile setae, 1 attenuate sensory seta; tibia IV, 13 tactile
setae, trichoboth; tarsus I, 11 plumose ventrals in 2 rows, 10 lateral

A Revision of the Mite Family Bdellidae 381

and 2 dorsal tactile setae, 2 attenuate and 2 blunt sensory setae, 1
attenuate peg, dorsoterminals solid, minutely pilose; tarsus II, 10
plumose ventrals, 7 lateral and 2 dorsal tactile setae, 2 blunt sensory
setae, 1 attenuate peg, dorsoterminals as in tarsus I; tarsus III, 12
plumose ventrals, 8 laterals, trichoboth, dorsoterminals as in tarsus
I; tarsus IV, 12 plumose ventrals, 6 laterals, trichoboth, 1 attenuate
sensory seta proximal to trichoboth, dorsoterminals as in tarsus I.

Male: Identical to female except in total length and genital
regfon; length, including gnathosoma, 880 [x. Genital region: Thir-
teen equal setae in linear arrangement on each genital plate; 10
paragenitals, anterior pair between coxae IV; genital discs small,
widely separated; genital aperture large, about one half the length
of the hysterosoma; amphioid sclerites (fig. 170) extending from
between coxae IV to near the ventral termination of anal cleft.

Holofype: Female, 8 miles north of Llera, Tamaulipas, Mexico,
July 19, 1954, W. T. Atyeo, beating pineapple epiphyte on mesquite.

Allotype: Male, Cuidad del Maiz, San Luis Potosi, Mexico; at
Laredo, Texas, July 10, 1945, T. P. Chapman, on orchid plants.

Paratijpes: One male, one female, 8 miles west of Antiguo More-
los, Tamaulipas, Mexico, July 21, 1954, W. T. Atyeo, under log; one
female, Mante, Tamaulipas, Mexico; at Laredo, Texas, March 28,
1945, C. D. Babb, on orchid plants.

Location of types: The holotype and two paratypes deposited in
the Snow Entomological Museum, the allotype and one paratype at
the United States National Museum.

Remarks: Except for the number of setae on the male genital
plates, which vary from twelve to thirteen pairs, and a slight varia-
tion in the number of tactile setae on the leg segments, characters
in this species appear to be constant. Total length varies from 902 \i
to 994 [L. This species is named lon<i,istriata to call attention to the
almost uninterrupted striae in the dorsal propodosomal pattern.
Drawings of the holotype.

Bdella distincta Baker and Balock

(Figs. 41, 79, 80, 141)

Bdella distincta Baker and Balock, 1944, Proc. Ent. Soc. Washington, vol. 46,
no. 7, p. 179.

This species is closely related to Bdella mexicana, but has dis-

tally branched dorsal hysterosomal setae, rather than long nude

or long plumose setae and has finely broken striae rather than

sparsely broken to continuous striae on the dorsal propodosoma.

382 The University Science Bulletin

Female: Color in life unknown. Body narrowly ovoid, strongly
constricted; length, including gnathosoma, 858 p,. Gnathosoma:
Length, 202 [i; palpus (fig. 41) short, distal two thirds of tibiotarsus
extending beyond hypostome; measurements: I, 10 p.; II plus III, 129
[i; IV, 29 [jl; V, 54 PL; des, 139 [j.; ves, 126 [>.. Chelicera normal, finely
striated, 187 [x in length; chela smooth, fixed digit attenuate, both
digits of equal length; distal seta extending to base of chela, proximal
seta extending half the distance to distal seta; setae inserted as in fig.
13. Hypostome nonstriated; ventral setae approximately equi-
distant, arranged in two longitudinal series, vh 1 between palpal
articulations; dorsal hypostomals slightly shorter than vh 1. Dorsal
propodosoma (fig. 141): Striae finally broken; lateral propodoso-
mals thickened, sparsely branched near apex, 51 [j. in length, median
propodosomals branched distally as in fig. 182, 53 \t. in length, eyes
separated by distance equal to diameter of anterior pair, space be-
tween eyes with transverse striae; distance between anterior sensilla,
95 \i.. Dorsal hysterosoma: Setae branched distally, as in fig. 182;
length of internal humeral, 49 p., approximately one half of first inter-
space; external humeral, 61 \i in length; sacrals in gently curving
transverse row, clunals in strongly curving transverse row. Anal
region: Border striae bending laterad; one pair of anal setae near
anterior termination of cleft; one pair of distally branched paranals
posterior to laterally directed striae; postanals branched, flanking
termination of cleft, slightly shorter than external clunals. Genital
region: Each genital plate with 8 small, equal setae in linear ar-
rangement: 9 pairs of paragenitals, anterior pair between coxae
IV; genital discs small, anterior pair one fourth the length of
vestibule from anterior end; two posterior pairs approximate, im-
mediately caudad of middle of vestibule; ovipositor with 12 sub-
apical, 6 postmedial setae. Legs ( figs. 79, 80 ) : Claws with one row
of small lateral rays each; coxae nonstriated; measurements: tibia I,
51 \).; tarsus I, 75 \i; tibia II, 32 [jl; tarsus II, 72 [i. Chaetotaxy: coxae
I-IV, 5, 4, 4, 3 tactile setae; trochanters I-IV, 1, 1, 2, 1 tactile setae;
basifemora I-IV, 8, 9, 7, 3 tactile setae; telofemora I-IV, each with 1
large, dorsal, branched seta plus 4, 5, 5, 3 tactile setae; genu I, 4 tac-
tiles, 2 attenuate sensory setae; genua II-IV, 5 tactile setae, 1 at-
tenuate sensory seta each; tibia I, 7 tactiles, 3 attenuate sensory
setae, 1 attenuate peg, trichoboth; tibia II, 8 tactiles, 1 attenuate and
1 blunt sensory seta; tibia III, 7 tactiles, 1 attenuate sensory seta;
tibia IV, 7 tactiles, trichoboth; tarsus I, 10 plumose ventrals in 2
rows, 5 lateral and 2 dorsal tactile setae, 2 attenuate and 2 blunt

A Revision of the Mite Family Bdellidae 383

sensory setae, 1 hollow, blunt peg, dt 1 scaly, dt 2, S minutely
plumose; tarsus II, 8 plumose ventrals, 5 lateral and 1 dorsal tactile
setae, 2 blunt sensory setae, 1 solid peg, dorsoterminals as in tarsus
I; tarsus III, 8 plumose ventrals, 4 laterals, trichoboth, dorsoter-
minals as in tarsus I; tarsus IV, 8 plumose ventrals, 4 laterals, tricho-
both, 1 attenuate sensory seta distal to trichoboth, dorsoterminals
as in tarsus I.
Male: Unknown.

Types: Female, China, at Washington, D. C, January 29, 1941,
on Bamhusa parvoriabilis tree; female, Hawaii, at Houston, Texas,
October 18, 1934, O. D. Morris, in pine cones. The specimen from
China is herewith designated lectotype by the present author.

Location of types: The United States National Museum, type no.
1463.

Remarks: The material examined by the present writer included
specimens with the following data: two females, Mexico, at Laredo,
Texas, August 24, 1945, I. A. Lane, on Croton cuttings; one female,
Guayama, Puerto Rico, October 20, 1941, G. N. Wolcott, on Ficiis
stahlii; one female, Philippines, at Honolulu, Hawaii, April 14, 1933,
on Saccolahiiim violaceum; two nymphs, Indonesia, at Washington,
D. C, April 19, 1954, H. Y. Goudeman, on cameHia cuttings.

The characters distinguishing this species are stable. The length,
including gnathosoma, varies from 688 jjl to 858 [jl, and the tactile
setae on the leg segments vary from one more, to one less, than the
numbers indicated in the redescription. Redescription and draw-
ings of the lectotype.

Bdella mexicana Baker and Balock

(Figs. 13,36,81,82)

Bdella mexicana Baker and Balock, 1944, Proc. Ent. Soc. Washington, vol.

46, no. 7, p. 181.
Bdella ivillisi Baker and Balock, 1944, Proc. Ent. Soc. Washington, vol. 46,

no. 7, p. 182 (new synonym).

This species is closely related to Bdella distincta, but has sparsely
broken to continuous striae in the dorsal propodosomal pattern,
rather than finely broken striae. Bdella mexicana is unique among
the known species of Bdella in that it lacks a trichoboth on tarsus IV.

Female: Color in life unknown. Body narrow, strongly con-
stricted; length, including gnathosoma, 560 [i. Gnathosoma: Length,
183 jx; palpus (fig. 36) short, distal third of tibiotarsus extending
beyond hypostome; measurements: 1, 11 [x; II plus III, 79 [i; IV, 20 [x;
V, 48 I).; des, 104 [jl; ves, 73 [jl. Chelicera ( fig. 13 ) inflated, finely stri-

384 The UNivERSiTi' Science Bulletin

ated, 148 pi in length; chela dentate, one small tooth on movable
digit, fixed digit attenuate, two thirds the length of movable digit;
distal seta extending to base of chela, proximal seta extending two
thirds the distance to distal seta, setae as figured. Hypostome non-
striated; ventral setae in two longitudinal series, vh 1 between pal-
pal articulations, setae in each row approximately equidistant; dor-
sal hypostomal setae equal in length to vh 1. Dorsal propodosoma:
Striae sparsely broken, pattern similar to BdeUa distincta (fig. 141);
lateral propodosomals nude, 30 [i in length; median propodosomals
nude, 34 [l in length; eyes separated by distance equal to one half
the diameter of anterior pair, space between eyes with longitudinal
striae; distance between anterior sensilla, 64 pi. Dorsal hysterosoma:
Setae minutely plumose; length of internal humeral, 45 pi, approxi-
mately one half of first interspace; external humeral, 61 pi in length;
sacrals and clunals in gently curving transverse rows. Anal region:
Border striae bending laterad; one pair of anal setae anterior, one
pair posterior of laterally directed striae; paranals wanting; post-
anals flanking termination of cleft, shorter than the clunal setae.
Genital region: Each genital plate with 8 small, equal setae in
linear arrangement inserted on a longitudinal, nonstriated area; 9
pairs of paragenitals, anterior pair between coxae IV, genital discs
small, anterior pair one fourth the length of the vestibule from
anterior end, posterior pairs approximate, immediately caudad of
middle; ovipositor with 12 subapical, 6 postmedial setae. Legs
(figs. 81, 82): Claws with one row of short, lateral rays each;
measurements: tibia I, 42 pi; tarsus I, 44 pi; tibia II, 26 \i; tarsus II,
32 pi. Chaetotaxy: coxae I-IV, 5, 4, 5, 2 tactile setae; trochanters
I-IV, 1, 1, 2, 1 tactile setae; basifemora I-IV, 8, 7, 7, 5 tactile setae;
telofemora I-IV, 5, 5, 5, 4 tactile setae; genua I-IV, 4 tactile setae,
one attenuate sensory seta each; tibia I, 6 tactile setae, 3 attenuate
sensory setae, trichoboth; tibia II, 5 tactile setae, one attenuate and
one blunt sensory seta; tibia III, 5 tactile setae, one attenuate
sensory seta; tibia IV, 5 tactile setae, trichoboth; tarsus I, 7 plu-
mose ventrals arranged in 2 rows, 5 lateral and 2 dorsal tactile
setae, 2 attenuate and 2 blunt sensory setae, one solid, attenuate
peg, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus II, 6 plumose
ventrals, 3 lateral and 1 dorsal tactile seta, 2 blunt sensory setae,
one solid, attenuate peg, dorsoterminals as in tarsus I; tarsus III,
6 plumose ventrals, 3 laterals, trichoboth, dorsoterminals as in tar-
sus I; tarsus IV, 6 plumose ventrals, 2 laterals, 1 attenuate sensory
seta, trichoboth wanting, dorsoterminals as in tarsus I.

A Revision of the Mite Family Bdellidae 385

Male: Identical to female; amphioid sclerites each with five equal
setae in gently curving longitudinal row on anterior half of plate,
three smaller setae in oblique row approximately two thirds the
length of the plate from the anterior end.

Type: Female, Valle del Bravo, Mexico, Mexico, March 4, 1943,
Donald Dodds, moss.

Location of type: The United States National Museum, type no.
1464.

Remarks: The type of Bdella willisi was also examined; this speci-
men, a male, was collected at Laguna de Zempoala, Morelos, Mex-
ico, January 31, 1943, by E. W. Baker, in moss. Ten additional
females were available and these included specimens from Mexico
(San Luis Potosi, Veracruz) and the United States (Texas, Kansas,
Illinois, Maryland, New York, Connecticut).

The dorsal hysterosomals are nude in the Mexican specimens,
but pilose in the individuals collected in the United States. Inter-
grades between these two conditions were not discovered, but pre-
sumably, they might occur in a limited area near the Texas-Mexican
border. In the study specimens, the number of tactile setae on
the leg segments, especially the basifemora, vary from two more
to one less than the number indicated in the description of the
holotype. The total length varies from 560 [i to 675 [x, and the other
measurements vary proportionately. Drawings of the mexicana
type.

Odontoscirinae Grandjean, 193S

The Odontoscirinae is characterized by six or seven pairs of
ventral hypostomal setae, five pairs of trichoboths, and the absence
of genital tracheae. The palpal tibiotarsus is cylindrical and
usually as long or longer than the pajpal basifemur. The end setae
are equal or subequal and usually shorter than the palpal tibio-
tarsus. Normal to inflated chelicerae bear one to many setae in-
serted on the proximal three fourths; the chelae have the movable
digits sickle- shaped (straight in Odontoscirus). Lateral propo-
dosomal setae may be present or absent; secondary apodcmes are
usually not developed, although large amounts of cuticular deposi-
tion may form large subcutaneous plates. The striation patterns on
the dorsum of the propodosoma are similar in all genera; four eyes
are lateral or posterolateral to the modified or unmodified posterior
pseudostigmata. The podocephalic canal is an internal tube
(Grandjean, 1938).

13—8920

386 The University Science Bulletin

Genus Odontoscirus Thor

Biscinis (Odontoscirus) Thor, 1913, Zool. Anz., vol. 42, no. 1, p. 29. (Type:
Bdclla virgulata Canestrini and Fanzago, 1876 [by original designation].)

The modifications of the gnathosoma are characteristic for this
distinctive genus. The chehcerae are inflated; the digits are ap-
proximately equal in length, and the movable digit is basically
straight and dentate. The lateral lips are much larger than the
same structures in other genera (except Cijfo). The lateral propo-
dosomal setae are wanting. The posterior pseudostigmatic organs
are unmodified and the posterior sensilla are longer than the median
propodosomal setae. Legs II are shorter than legs I.

Key to the Species of Odontoscirus

1. Small; chelicerae and gnathosoma striated; palpal tibiotarsus with
4 tactile setae, 1 attenuate sensory seta, and 2 long apical setae

iota p. 386
Large; only gnathosomal base striated; palpal tibiotarsus with 6
tactile setae, 1 attenuate sensory seta, and 2 long apical setae

alpinus p. 388

Odontoscirus iota, sp. nov.
(Figs. 9, 16, 17, 44, 83, 84, 154)

This species is related to Odontoscirus virgulatus (Canestrini and
Fanzago), 1876, but is about 900 [i to 1100 [j. rather than 1100 ;;. to
2000 \L in total length. The palpal tibiotarsus of O. iota is approxi-
mately 120 [I in length, as compared to the same structure of O.
virgulutus, which is approximately 250 ^ long.

Female: Color unknown. Body narrowly ovoid; length, includ-
ing gnathosoma, 1100 [x. Gnathosoma: Length, 264 |x; palpus (fig.
44) relatively short, tibiotarsus extending beyond hypostome;
measurements: I, 15 pi; II, 158 [}.; Ill, 26 \>.; IV, 22 [k; V, 117 ii; des,
145 [jl; ves, 150 [j,. Chelicera inflated, finely striated, 255 pi, in length;
chela (figs. 9, 17) dentate, movable digit with 4 small teeth, fixed
digit with small subapical tooth, digits of same length; setae inserted
as in fig. 16. Gnathosoma striated; ventral setae in two longitu-
dinal series; vh 1-5 approximately equidistant, vh 6 medial between
vh 5 and apices of lateral lips, vh 2 between palpal articulations;
vh 1 caudad; dorsal hypostomal setae equal in length to vh 3. Dor-
sal propodosoma (fig. 154): Striae finely broken; median propodo-
somals nude, 80 [i in length; eyes separated by a distance equal to
two diameters of anterior pair; distance between anterior sensilla,
66 [).. Dorsal hijsterosoma: Setae nude; length of internal humeral,
65 ]}., approximately one half of first interspace; external humeral,

A Revision of the Mite Family Bdellidae 387

82 [x in length; sacrals in gently curving transverse row; clunals in
subrectangular arrangement. Anal region: Border striae parallel;
one pair of anal setae; paranals wanting; postanals flanking termi-
nation of cleft, shorter than clunal setae. Genital region: Each gen-
ital plate with 7 equal genital setae in linear arrangement; 3 pairs
of paragenitals caudad of anterior termination of aperture; genital
discs small, anterior pair slightly removed from proximate posterior
pairs; ovipositor with 16 subapical, 6 postmedial setae. Legs
(figs. 83, 84): Claws with 4-5 lateral rays each, minute ravs want-
ing; measurements: tibia I, 68 [;.; tarsus I, 155 [j.; tibia II, 60 [j.; tarsus
II, 143 ]x. Chaetotaxy: coxae I-IV, 5, 2, 4, 2 tactile setae; trochanters
I-IV, 1 tactile seta each; basifemora I-IV, 11, 11, 7, 3 tactiles, 5 at-
tenuate sensory setae; genu I, 6 tactiles, 5 attenuate sensory setae;
genu II, 6 tactiles, 2 attenuate sens^ory setae; genua III-IV, 5 tactiles
and 1 attenuate sensory seta each; tibia I, 10 tactiles, 4 attenuate and
1 blunt sensory seta, 1 peg, trichoboth; tibia II, 8 tactiles, 1 attenuate
and 1 blunt sensory seta, trichoboth; tibia III, 10 tactiles, 1 attenuate
sensory seta; tibia IV, 9 tactiles, trichoboth; tarsus I, 14 plumose
ventrals arranged in two rows, 6 lateral and 2 dorsal tactile setae, 1
attenuate and 3 blunt sensory setae, 1 peg, dorsal terminals, solid,
plumose; tarsus II, ventrals, laterals, and dorsoterminals as in tarsus
I, 1 dorsal tactile seta, 1 attenuate and 2 blunt sensory setae, 1 peg;
tarsus III, ventrals, laterals, and dorsoterminals as in tarsus I, tricho-
both; tarsus IV, ventrals and laterals as in tarsus I, trichoboth, dt 1
reduced to 1 solid, plumose seta, dt 2, 3 solid, plumose.

Male: Identical to female; slightly longer than female; amphioid
sclerites with 3, 1, 3, 2 setae each.

Holotype: Female, near Oakland, California (in hills), Febru-
ary 12, 1956, N. A. Walker, redwood.

Allotype: Male, Naperville, Illinois, January 24, 1931, C. C.
Compton, in mushrooms.

Paratypes: Two females, same data as holotype.

Location of types: Holotype and paratypes deposited in the
Snow Entomological Museum; the allotype deposited at the United
States National Museum.

Remarks: Other than slight size diflFerences, the only notable
variation occurs in one female, which has ten pairs of setae on the
genital plates, rather than seven pairs. This species is named
iota to denote its small size when compared to the other described
species of Odontoscirus. Drawings of the holotype.

388 The University Science Bulletin

Odontoscirus alpinus, sp. nov.

(Figs. 15, 43, 85, 86)

This species is closely related to O. virgulatus (Canestrini and
Fanzago), 1876, but has three to four teeth on the movable digit
of the chelicera instead of four to five teeth. Also, the palpal
tibiotarsus of alpinus has two setae on the proximal half, whereas
virgulatus has all of the setae restricted to the distal half of that
segment.

Female: Color in life red with dark blue to black blotches. Body
ovoid, weakly constricted; length, including gnathosoma, 1990 \i.
Gnathosoma: Length, 533 pi; palpus (fig. 43) with tibiotarsus ex-
tending beyond hypostome; measurements: I, 12 [i; II, 189 [i; III,
26 [t.; IV, 24 [k; V, 164 ii; des, 238 [jl; ves, 238 ij.. Chelicera normal,
reticulated, 476 [x in length; digits equal in length, movable digit
with 4-5 teeth, fixed digit with 1 subapical tooth; setae inserted
medially (fig. 15). Cnathosomal base striated, buccal cone non-
striated; ventral setae in two longitudinal series, vh 1-4 equidistant,
interval between vh 4-5 approximately twice the distance between
vh 2 and 3, vh 6 near the termination of buccal cone, vh 2-4 inserted
on the proximal half of the cone, vh 1 on base; dorsal hypostomals
equal in length to vh 3. Dorsal propodosoma: Striae finely broken;
pattern similar to that of Odontoscirus iota (fig. 154); median pro-
podosomals nude, approximately 100 pi in length; eyes separated
by distance equal to four times the diameter of the anterior pair;
distance between anterior sensilla, 78 [x. Dorsal hysterosoma: Setae
finely plumose; length of internal humeral, 104 [;., approximately
one third of first interspace; external humeral, 109 [jl in length;
sacrals in gently curving transverse row, clunals in subrectangular
arrangement. Anal region: Border striae parallel; one pair of anal
setae; paranals wanting; postanals flanking termination of cleft,
shorter than clunal setae. Genital region: Each genital plate with
7 equal attenuate setae in linear arrangement; 3 pairs of para-
genitals ; genital discs of medium size, proximate, in central area
of vestibule; ovipositor with 16 subapical, 6 postmedial setae. Legs
( figs. 85, 86 ) : Claws with 6 lateral rays each, minute rays
wanting; measurements: tibia I, 158 [x; tarsus I, 311 |j.; tibia 11,
151 [i; tarsus II, 298 [x. Chaetotaxy: coxae I-IV, 5, 3, 4, 2 tactile
setae; trochanters I-IV, 1 tactile seta each; basifemora I-IV, 11, 13,
9, 3 tactile setae; telofemora I-IV, 7, 8, 7, 7 tactile setae; genu I, 6
tactiles, 6 attenuate sensory setae; genua II-IV, 6 tactile and 3
attenuate and two blunt sensory setae, 1 peg, dt 1, solid, nude.

A Revision of the Mite Family Bdellidae 389

setae, 1 peg, trichoboth; tibia II, 10 tactiles, 1 attenuate and 1 blunt
sensory seta, trichoboth; tibia III, 12 tactiles, 1 attenuate sensory
seta; tibia IV, 13 tactiles, trichoboth; tarsus I, 22 plumose ventrals
arranged in two rows, 10 lateral and 1 dorsal tactile seta, two
attenuate and two blunt sensory setae, 1 peg, dt 1, solid, nude,
dt 2, 3 solid, plumose; tarsus II similar to tarsus I except lacking 1
lateral tactile seta and 1 attenuate sensory seta; tarsus III, 22
plumose ventrals, 9 laterals, dorsoterminals as in leg I, trichoboth;
tarsus IV, similar to tarsus III, but dt 1 reduced to 1 solid, nude seta.

Male: Unknown.

Holotype: Female, Piegan Pass, Glacier National Park, Montana,
August 23, 1953, collector: Levi, altitude 7,900 ft.

Parafype: One female, same data as holotype.

Location of types: The holotype and paratype deposited in the
Snow Entomological Museum.

Remarks: These specimens agree in general size with O. vir^ii-
latus, but the critical features for determination are lacking in the
description of the European species. The paratype female is slightly
smaller than the holotype, total length, 1030 \l, but the other features
are constant. This new species is named alpinus because the only
known specimens were collected at high altitudes. Drawings of the
holotype.

Genus Neomolg,us Oudemans

Neomolgus Oudemans, 1937, Kritisch Historisch Overzicht der Acaro^ogie,

Leiden, vol. 3, part C, p. 1229. (Type: Acartis littoralis L., 1758 [by

orij^inal designation].)
Molffus Trouessart, 1894 (non Dujardin 1842, Murray 1877, Halacaridae) Jour.

Ant. Physio'., vol. 30, p. 117. (Type: Molgns sanguineus Trouessart, 1894

[= Acarns littoralis L.] [by original designation].)
Molgiis (Hoplornolgiis) Berlese, 1923, Redia. vol. 15, p. 237 (new synonym).

(Type: Bdella capillata Berlese, 1891 [non Kramer, 1881] [by original

designation].)

This genus, which is related to Bdellodes, is characterized as
having more than two setae on each chclicera. The chelicerae are
normal; the movable digits are smooth or flattened distally on the
inner surface, and are slightly longer than the fixed digits. The
lateral propodosomal setae are wanting. The posterior pseudostig-
mata are unmodified and the posterior sensilla are longer than the
median propodosomal setae. Legs II are shorter than legs I.

The subgenus Hoplomolgus was erected for species of Neomolgus
(z= Molf^us) with distinct, reniform dorsal shields. These shields,
formed by a large amount of cuticular deposition, are considered
to be extreme modifications of a continuous character, /. e., cuticular
deposition.

390 The University Science Bulletin

Key to the Species of Neomolgus

1. Chelicera with 9-10 setae; palpal basifemur with 6-8 setae

littoralis p. 390
Chelicera with 6-8 setae; palpal basifemur with 3-4 setae

mutabilis p. 392

Neomolgus littoralis ( L. )
(Figs. 29, 56, 107, 108)
Acarus littoralis Linnaeus, 1758, Syst. Nat., ed. 10, p. 618.

This species appears to be closely related to N. capillatus
(Kramer), 1881. Although of comparable size, N. littoralis has fif-
teen to twenty-four setae on the palpal tibiotarsus rather than nine
to eleven, and has long rather than short dorsal hysterosomal setae.

Female: Color in life dark red with irregular dark blue blotches.
Body robust, weakly constricted; length, including gnathosoma,
2200 [J. (2060-2345 [x). Gnathosoma: Length, 575 [;. (518-660 [jl);
palpus (fig. 56) with genu two times longer than telofemur; meas-
urements: I, 31 PL (29-34 [l); II, 422 fx (315-476 [x); III, 94 i,. (65-
119 [x); IV, 168 [x (122-185 [l);V, 400 [l (325-425 [x); des, 170 ;x
146-170 pl); ves, 153 jx (119-158 [l). Chelicera (fig. 29) normal
nonstriated, 568 [x (511-617 p.) in length, with 10 (9-10) setae;
movable digit with inner surface flattened apically, fixed digit ex-
tending to proximal termination of flattened surface of movable
digit. Gnathosomal base striated, buccal cone nonstriated; ventral
setae form two longitudinal rows, vh 1-5 on proximal third of cone,
vh 6 medial between apex and vh 5; dorsal hypostomal setae as long
as vh 1. Dorsal propodosoma: Striae finely broken; pattern similar
to fig. 163, median propodosomals nude (to finely plumose), 235 ;x
(212-272 [x) in length; eyes separated by distance equal to 3
diameters of anterior pair, interval between eyes with transverse
striae; distance between anterior sensilla, 119 -jX (103-122 [x). Dorsal
hysterosoma: Setae finely plumose; length of internal humeral,
174 [X (141-174 [x), approximately half of first interspace; external
humeral, 243 \i (170-280 [x) in length; sacrals in gently curving
transverse row, clunals in strongly curving transverse row. Anal
region: Border striae parallel; anal setae wanting; three pairs of
paranals; postanals flanking termination of cleft, shorter than clunals.
Genital region: Each genital plate with 12, approximately equal,
attenuate setae in irregular linear arrangement; 7 paragenitals, an-
terior pair at level of anterior termination of aperture; genital
discs large, anterior pair exceeding twice the length of the median
pair; ovipositor with 20 subapical setae. Legs (figs. 107, 108):
Claws with 7-8 lateral rays each; measurements: tibia I, 247 [x

A Revision of the Mite Family Bdellidae 391

(192-272 ii); tarsus I, 388 [jl (308-425 [x); tibia II, 244 [x (187-260 [jl);
tarsus II, 383 pt (306-418 ix). Chaetotaxy: coxae I-IV, 11 (8-12), II
(9-11), 13 (12), 12 (7-12) tactile setae; trochanters I-IV, 2, 2, 3 (2),
3(4) tactile setae; basifemora I-IV, 13 (12-18), 13 (13-16), 11 (10),
8 (9) tactile setae; telofemora I-IV, 7 (7-11), 7 (8), 7 (8), 6 (7)
tactile setae; genu I, 6 (6-9) tactile setae, 14 (9-14) attenuate sen-
sory setae; genu II, 6 (6-8) tactile setae, 10 (5-11) attenuate sensory
setae; genu III, 6 ( 5-7 ) tactile setae, 8 ( 5-8 ) attenuate sensory setae;
genu IV, 8(9) tactile setae, 5 (3-5) attenuate sensory setae; tibia I,
12 (11 18) tactile setae, 20 (13-23) attenuate sensory setae, 1 peg,
trichoboth; tibia II, 12 (10-16) tactile setae, 13 (10-20) attenuate
sensory setae, 1 blunt sensory seta, trichoboth; tibia III, 14 (14-16)
tactile setae, 7 (4-9) attenuate sensory setae; tibia IV, 13 (13-14)
tactile setae, 3 ( 1-4 ) attenuate sensory setae, trichoboth ( may be
absent); tarsus I, 28 (27-34) ventrals, distal pairs plumose, arranged
in 4 rows, proximal pairs nude, in 2 rows, 10 ( 13 ) lateral, 2 dorso-
lateral and 2 dorsal tactile setae, 7 (5-7) attenuate sensory setae
and 2 blunt sensory setae, 1 peg, dt 1 solid, nude, dt 2, 3 solid,
coarsely plumose; tarsus II, identical to leg I, except lacking 1 dorso-
lateral tactile seta, dt 2, solid, nude; tarsus III, ventrals and laterals
as in leg I, 3 attenuate sensory setae, trichoboth, dorsoterminals as
in leg II; tarsus IV, similar to leg III, except 2 attenuate sensory
setae, dt 1 reduced to one seta.

Male: Identical to female; amphioid sclerites with 3, 4, 2 setae
each.

Type: Europe.

Location of type: Unknown.

Remarks: Over one hundred and fifty specimens were examined,
which included collections from: Alaska, Hudson Bay area, Green-
land, Iceland, Spitsbergen, Bering Island, Russian Lapland (Kola-
Hafvon), Nova Zembla (Matochkin Strait), and Finland.

The specimens from Matochkin Strait, four males, are of special
interest. Typical in other characteristics, each of these mites lacks
the trichoboth on tibia IV, a long tactile seta is inserted at the posi-
tion normally occupied by the trichoboth.

Thor (1931) reports a greater variation in size and chaetotaxy
in this species than indicated in the present study. Total length
varies from 1500 jj. to 3500 ii and other measurements vary accord-
ingly. The setae of the palpus may vary as follows: II, 6-8; III, 1;
IV, 5-7; V, 15-24 tactile setae. The redescription and illustrations
are based on a female collected at Sanders Island, Greenland, Au-
gust 4, 1952, J. M. Anderson, Ex: beach.

392 The University Science Bulletin

Neomolgtis mutabilis, sp. nov.
(Figs. 28, 55, HI, 112, 163)

This species is closely related to Neomolgus pallipes ( Koch ) , 1879,
but can be distinguished by the number of setae on the palpal
basifemur and chelicera, respectively, 3-4 rather than 5-6, and 6-8
rather than 8-10.

Female: Color in life unknown. Body ovoid, weakly constricted;
length, including gnathosoma, 1660 [l. Gnathosoma: Length, 413 [jl;
palpus (fig. 55) with tibiotarsus approximately equal in length to
basifemur, genu equal to telofemur; measurements: I, 23 (x; II,
216 [x; III, 34 [jl; IV, 44 ;j.; V, 235 [x; des, 136 pi; ves, 124ti. Chelicera
(fig. 28) normal, nonstriated with 7 setae, 371 [x in length; movable
digit distally flattened on inner surface, fixed digit extending to
proximal termination of flattened surface of movable digit. Gnath-
osomal base striated, buccal cone nonstriated; ventral setae form
two longitudinal rows, vh 1-5 approximate, equidistant, vh 6 medial
between apex and vh 5; dorsal hypostomal setae equal in length to
vh 2. Dorsal propodosoma (fig. 163) : Striae finely broken; median
propodosomals plumose, 65 [i in length; eyes separated by distance
equal to 3 diameters of anterior pair; interval between eyes with
transverse striae; distance between anterior sensilla, 65 \).. Dorsal
htjsterosoma: Setae plumose; length of internal humeral, 43 {x, ap-
proximately one fourth of first interspace; length of external humeral,
53 [X; sacrals in gently curving transverse row, clunals in subrectan-
gular arrangement. Anal region: Border striae parallel, anals want-
ing; one pair of paranals; postanals flanking termination of cleft,
equal in length to clunals. Genital region: Each genital plate with
8 progressively shorter setae in linear arrangement; five pairs of
paragenitals, anterior pair between coxae IV; genital discs large,
equidistant, in anterior three fourths of vestibule; ovipositor with
16 subapical and 4 postmedial setae. Legs (figs. Ill, 112): Claws
each with 4-6 lateral rays, one row of minute rays reduced in num-
ber; measurements: tibia I, 136 [x; tarsus I, 197 [x; tibia II, 129 [i;
tarsus II, 187 \i. Chaetotaxy: coxae I-IV, 5, 5, 5, 3 tactile setae;
trochanters I-IV, each with one tactile seta; basifemora I-IV, 10, 9,
7, 4 tactile setae; telofemora I-IV, 8, 7, 6, 6 tactile setae; genu I, 6
tactiles, 12 attenuate sensory setae; genu II, 6 tactiles, 5 attenuate
sensory setae; genu III, 5 tactiles, 5 attenuate sensory setae; genu
IV, 6 tactiles, 4 attenuate sensory setae; tibia I, 12 tactiles, 13
attenuate sensory setae, 1 peg, trichoboth; tibia II, 12 tactiles, 5
attenuate sensory setae, 1 blunt sensory seta, trichoboth; tibia III,

A Revision of the Mite Family Bdellidae 393

11 tactiles, 3 attenuate sensory setae; tibia IV, 11 tactiles, 3 attenuate
sensory setae, trichoboth; tarsus I, 15 ventrals arranged in 2 rows,
distal 11 plumose, 7 lateral and 2 dorsal tactile setae, 4 attenuate
and 2 blunt sensory setae, 1 peg, dt 1 anterior to dt 2, solid, nude,
dt 3 between dt 2 solid, plumose; tarsus II, 15 ventrals, distal 11
plumose, 6 lateral and 2 dorsal tactiles, 2 blunt sensory setae, 1 peg,
dt 1, 2 solid, nude, dt 3 solid, plumose; tarsus III, ventrals and dorso-
terminals as in leg II, 6 laterals, trichoboth; tarsus IV, 14 ventrals,
distal 8 plumose, 5 laterals, trichoboth, df 1 reduced to 1 solid nude
seta, dt 2, 3 as in leg II.

Male: Closely resembles the female except in size, number of
attenuate sensory setae and dorsal tactile setae on the legs and in
the genital region. Length, including gnathosoma, 1320 pi, other
measurements decrease accordingly; number of attenuate sensory
setae: genua I-IV, 10, 6, 5, 4; tibiae I-IV, 11, 7, 3, 4; tarsus I with 3
dorsal tactile setae; each genital plate with 7 setae; amphioid
sclerites with 3, 4, 2 setae each.

Holotype: Male, 2 miles south of Galena, Cherokee Co., Kansas,
April 8, 1955, Ronald B. Winslow, under board.

Allotype: Female, 2 miles south of Galena, Cherokee Co., Kansas,
April 8, 1955, W. T. Atyeo, under stone.

Paratypes: Four males, same data as holotype; one male, one
female, same data as allotype; one male, one female, Univ. Kansas
campus, Lawrence, Douglas Co., Kansas, April 22, 1952, R. E. Beer,
under rock; three males, Univ. Kansas campus, collected re-
spectively April 4, April 6, August 26, 1955, D. S. Narayan, under
rocks; one male, Spring Hill, Johnson Co., Kansas, April 7, 1956,
D. S. Lang, under rock; one male, 5 miles south of Avila Camacho,
Distrito Federal, Mexico, July 3, 1956, R. E. Beer, moss (Berlese
funnel extraction); one male, Mt. Popocatepetl, Mexico, Mexico,
August 18, 1954, W. T. Atyeo, beating Hchens on trees, alt., 13,000
feet; one male, Denison, Texas, February 19, 1938, L. D. Christen-
son, in soil; one male, 13 miles east of Toluca, Mexico, Mexico,
August 17, 1954, W. T. At)^eo, under rocks.

Location of types: The holotype, allotype and thirteen paraty|-)es
are deposited in the Snow Entomological Museum; one paratxqio
deposited at each of the following: the United States National
Museum, the British Museum (Natural History) and the South
Australian Museum.

Remarks: The chaetotaxy of the gnathosoma, genital plates and
appendages exhibit considerable variation. The Mexican specimens

394 The Universiiy Science Bulletin

have one seta on the proximal half of the palpal tibiotarsus; speci-
mens from the United States have two to three setae in this area.
Sporadically, the palpal basifemora have an extra median seta, the
chelicerae have six to eight setae, and the external genital flaps
have seven to nine setae. The number of attenuate sensory setae on
the legs varies as follows: telofemur I, 0-2; genua I-IV, 9-13, 5-6,
3-5, 2-4; tibiae I-IV, 6-13, 4-7, 2-5, 2-4. The attenuate sensory setae
on telofemur I are of interest; this is one of the few species in which
special sensory setae occur on the segments proximal to the genu.
Total length varies from 944 [t. to 1500 [>.. This species is named
mtitahilis to indicate the extreme variability. Drawings are of the
holotype.

Genus Thoribdella Grandjean

Thoribdella Grandjean, 1938, Ann. Soc. Ent. France, vol. 107, p. 4. (Type:
Biscirus meridionalis Thor, 1931 [by original designation].)

This genus, which is probably closely related to Bdellodes, is
distinguished by the deep goblet-shaped posterior pseudostigmatic
organs and usually by the extreme reduction of the posterior sen-
silla. The chelicerae are normal and each has one or two setae.
The inner surfaces of the movable digits are usually flattened dis-
tally, and are usually longer than the fixed digits. The lateral
propodosomal setae may be present or absent. Legs II are as
long or longer than legs I,

Key to the Species of Thoribdella

1. Posterior sensilla longer than median propodosomal setae; distal

cheliceral seta long, spinelike spinosa p. 395

Posterior sensilla much shorter than median propodosomal setae;

distal chehceral seta not spinelike 2

2. Posterior sensilla removed from median propodosomal setae; palpal

genu twice as long as palpal telofemur calif arnica p. 396

Posterior sensilla approximate to median propodosomal setae; pal-
pal genu and telofemur of approximately equal lengths 3

3. Lateral propodosomal setae present insolita p. 398

Lateral propodosomal setae absent 4

4. Palpal genu with four setae; palpal tibiotarsus shorter than basi-

femur; internal humeral as long as first interspace communis p. 399

Palpal genu with tliree setae; palpal tibiotarsus as long as or longer

than basifemur; internal humeral one third to one half of first inter-
space 5

5. Palpal basifemur three fourths the length of the tibiotarsus; pos-
terior sensilla thickened, truncated truncata p. 402

Palpal basifemur and tibiotarsus of approximately equal lengths;
posterior sensilla setaceous 6

A Revision of the Mite Family Bdellidae 395

6. Palpal tibiotarsus with seven setae (including end setae) restricted

to distal half; cheliceral setae approximate meridionalis p. 404

Palpal tibiotarsus with twelve setae (including end setae) not
restricted to distal half; cheliceral setae distant simplex p. 405

Thoribdella spinosa sp. nov.
(Figs. 27, 54, 109, 110, 143)

This unique species has the distal cheHceral seta modified into
a long, thickened, spinelike structure which is bifid at the base,
thus forming a short basal branch. The posterior sensilla are dis-
tant from, and longer than the median propodosomal setae.

Female: Color unknown. Body ovoid, weakly constricted;
length, including gnathosoma, 1000 \h. Gnathosomu: Length, 391 [x;
palpus (fig. 54) short, distal five sixths of tibiotarsus extending
beyond hypostome; measurements: I, 14 [jl; II, 182 [x; III, 36 [jl; IV,
49 [jl; V, 150 [x; des, 185 [jl; ves, 160 [x. Chelicera (fig. 27) normal,
nonstriated, 306 [x in length; chela smooth; distal seta large with
short basal branch, proximal seta fine, approximate. Gnathosomal
base, proximal third of buccal cone striated, distal two thirds non-
striated; vh 1 inserted between palpal articulations, vh 1-5 equidis-
tant, arranged in 2 longitudinal series, vh 6, medial to series, mid-
way between vh 5 and apex; dorsal hypostomal setae shorter than
vh 1. Dorsal propodnsoma: Striae finely broken, pattern similar to
fig. 142; median propodosomals nude (fig. 143), 54 ij. in length;
posterior sensilla fine, 85 [x in length, distant from median propo-
dosomals; eyes separated by distance equal to two and one half
diameters of anterior pair; distance between anterior sensilla, 73 [x.
Dorsal htjsterosoma: Setae nude; length of internal humeral, 66 |x,
approximately one half of first interspace; external humeral, 66 [x in
length; sacrals in gently curving row, clunals in subrectangular
arrangement. Anal region: Border striae parallel; 1 pair of anal
setae, paranals wanting; postanals flanking termination of cleft,
shorter than clunals. Genital region: Each genital plate with 7
equal setae in linear arrangement; 2 pairs of paragenitals; ante-
rior 2 pairs of genital discs large, proximate, posterior pair small,
near caudal termination of vestibule; ovipositor with 14 subapical,
6 postmedial setae. Legs (figs. 109, 110): Claws with 5-6 lateral
rays each, row of minute rays wanting; measurements: tibia I, 71 [x;
tarsus I, 214 \).; tibia II, 73 jj.; tarsus II, 216 [x. Chaetotaxy: coxae
I-IV, 4, 2, 2, 2 tactile setae; trochanters I-IV, 1 tactile seta each;
basifemora I-IV, 11, 8, 5, 3 tactile setae; telofemora I-IV, 6, 6, 5, 4
tactile setae; genu I, 4 tactiles, 6 attenuate sensory setae; genu II,

396 The University Science Bulletin

4 tactiles, 3 attenuate sensory setae; genu III, 3 tactiles, 1 attenuate
sensory seta; genu IV, 4 tactiles, 1 attenuate sensory seta; tibia I,
7 tactiles, 3 attenuate sensory setae, trichoboth; tibia II, 7 tactiles,
1 attenuate and 1 blunt sensory seta, trichoboth; tibia III, 7 tac-
tiles, 1 attenuate sensory seta; tibia IV, 6 tactiles, trichoboth; tarsus
I, 14 ventrals arranged in 2 rows, distal 12 plumose, 7 lateral and no
dorsal tactile setae, 2 attenuate and 2 blunt sensory setae, 1 peg,
dt 1 solid, nude, dt 2, S solid, plumose; tarsus II, similar to tarsus
I but lacking attenuate sensory setae; tarsus III, ventrals and dor-
soterminals as in tarsus I, 2 laterals, 4 large dorsolaterals, tricho-
both; tarsus IV, ventrals as in tarsus I, 4 large dorsolaterals, tricho-
both, dt 1 reduced to one solid, nude seta, dt 2, 3 solid, plumose.

Male: Unknown.

Holotype: Female, Sierra del Rosario: El Rangel, Cuba, Janu-
ary, 1938, Alex Bierig, en hojarasca.

Location of type: The Snow Entomological Museum.

Remarks: The unique structure of the larger cheliceral seta and
the relative positions and lengths of the posterior sensilla and me-
dian propodosomal setae, warrants the naming of this species, even
though based on a single specimen. The name spinosa is chosen
to emphasize the peculiar form of the cheliceral seta. Drawings
of the holotype.

Thoribdella calif ornica (Banks), new combination

(Figs. 25, 52, 105, 106, 144)

Bdella californica Banks, 1904, Proc. California Acad. Sci., ser. 3, vol. 3, no.

13, p. 366.
Bdella mogna Ewing, 1913, J. Ent. Zool., vol. 5, p. 123 (new synonym).

This species is related to T. simplex, sp. nov., but has the palpal
genu twice as long as the telofemur rather than the two segments
of approximately equal length.

Male: Color in life red, with dark blue areas. Bodv ovoid,
weakly constricted; length, including gnathosoma, 1880 [i. Gnatho-
soma: Length, 497 [x; palpus (fig. 52) with genu approximately
twice the length of the telofemur; measurements: I, 24 [x; II, 306 [x;
III, 78 tx; IV, 122 !x; V, 325 ix; des, 221 \).- ves, 146 !x. Chelicera (fig.
25 ) normal, nonstriated, 483 [x in length; inner face of movable digit
flattened distally; setae as figured. Gnathosomal base striated,
buccal cone nonstriated; vh 1-6 in linear series, vh 1-5 progressively
more distant, vh 6 midway between vh 5 and base of lateral lips;
dorsal hypostomal setae thickened, half the length of vh 1. Dorsal

A Revision of the Mite Family Bdelxidae 397

propodosotrui: Striae finely broken, pattern similar to fig. 142;
median propodosomals (fig. 144) with almost parallel margins,
331 |x in length; posterior sensilla fine, distant from median propo-
dosomals, 51 [I in length; eyes separated by distance equal to one
and one half diameters of anterior pair; distance between anterior
sensilla, 100 [i. Dorsal hysterosoma: Setae minutely pilose; length
of internal humeral about 153 pi, approximately half of first inter-
space; external humerals broken, length unknown; sacrals and
clunals respectively in gently and strongly curving transverse rows.
Anal region: Border striae parallel; anal setae wanting; one pair
of paranals; postanals flanking termination of cleft, shorter than
clunals. Genital region: Each genital plate with 10 unequal setae
in linear arrangement, anterior 7 longer than posterior 3 setae; 3
pairs of paragenitals; genital discs of medium size, equidistant;
amphioid sclerites with 3, 1, 4, 1 setae each. Legs (figs. 105, 106):
Claws with 5-7 lateral rays each, row of minute rays wanting;
measurements: tibia I, 184 [x; tarsus I, 318 p.; tibia II, 201 li.; tarsus
II, 331 [X. Chaetotaxy: coxae I-IV, 4, 3, 4, 3 tactile setae; trochan-
ters I-IV, 1 tactile seta each; basifemora I-IV, 13, 13, 8, 4 tactile
setae; telofemora I-IV, 8, 8, 6, 6 tactile setae; genu I, 7 tactiles, 5 at-
tenuate sensory setae; genu II, 6 tactiles, 1 attenuate sensory seta;
genua III-IV, 6 tactiles and 2 attenuate sensory setae each; tibia I,
14 tactiles, 6 attenuate sensory setae, 1 attenuate-blunt sensory seta,
1 peg, trichoboth; tibia II, 11 tactiles, 2 attenuate sensory setae,
1 blunt sensory seta, trichoboth; tibia III, 12 tactiles, 1 attenuate
sensory seta; tibia IV, 12 tactiles, trichoboth; tarsus I, approxi-
mately 60 ventrals, distal two thirds plumose and arranged in 5-6
irregular rows, proximal third nude and arranged in 4 irregular
rows, 9 lateral and 2 dorsal tactile setae, 2 attenuate and 2 blunt
sensory setae, 1 peg, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus
II, approximately 50 ventrals arranged as in tarsus I, 10 lateral tac-
tiles, 1 attenuate and 2 blunt sensory setae, 1 peg, dorsoterminals as
in tarsus I; tarsus III, 40 ventrals, proximal half nude and arranged
in 2 irregular rows, distal half plumose, in 4 irregular rows, 9 lat-
erals, trichoboth, dorsoterminals similar to tarsus I; tarsus IV, ven-
trals and laterals as in tarsus III, trichoboth, dt 1 reduced to 1
solid, nude seta, dt 2, 3 solid, plumose.

Female (incomplete specimen): Similar to male; genital plates
with 8 pairs of setae; ovipositor with 22 setae of indeterminable
position.

Type: Claremont, Los Angeles Co., California.

398 The University Science Bulletin

Location of type: Unknown.

Remarks: Two cotypes of Thoribdella nwgna (Banks), 1913
{^= Bdella magna), one from the United States National Museum
and one from the Illinois Natural History Survey, were available
for examination. As these two specimens, both females, were in-
complete, it is impossible to determine whether or not any differ-
ences exist between the sexes other than the number of genital setae.

This species illustrates a distinct modification of the dorsal pro-
podosomal setae. The reduced posterior sensilla are inserted in
simple, straight-sided cups, and are distant from the long, almost
parallel-margined median propodosomals. Drawings and redescrip-
tion are based on a male collected at Oakland, Alameda Co., Cali-
fornia, January 30, 1953, W. C. Bentinck, habitat unknown.

Thoribdella insolita, sp. nov.
(Figs. 26, 53, 99, 100, 148)

This unique species has four pairs of dorsal propodosomal setae
and seven pairs of ventral hypostomal setae. Although reminiscent
of Octohdellodes species, insolita has the posterior sensilla modified
to structures typical of the genus Thoribdella. At the present time,
the lateral propodosomal setae are sufficient to distinguish this new
species.

Female: Color in life unknown. Length, including gnathosoma,
1540 [}.. Gnathosoma: Length, 468 [l; palpus (fig. 53) short, only
half of the tibiotarsus extending beyond gnathosoma; measurements:
I, 25 [k; II, 209 [i; III, 46 [x; IV, 53 [jl; V, 190 ;;.; des, 277 [l; ves,
216 [JL. Chelicera (fig. 26) normal, nonstriated, 442 [i in length;
movable digit with inner surface distally flattened; fixed digit with
preapical tooth, setae as figured. Gnathosoma striated to level of
vh 6, distally nonstriated; vh 1-6 approximately equidistant, in 2
longitudinal series; dorsal hypostomals about half the length of
vh 1. Dorsal propodosoma: Striae finely broken, pattern similar to
fig. 142, except a pair of lateral propodosomal setae inserted anterad
of posterior sensilla, approximately one fourth the interval between
posterior sensilla and anterior sensilla; lateral propodosomals nude,
146 \). in length; median propodosomals (fig. 148) with basal flange,
442 pt, in length; posterior sensilla approximate to median propo-
dosomals, 36 [X in length; eyes separated by distance equal to one
and one half diameters of anterior pair; distance between anterior
sensilla, 91 \i. Dorsal htjsterosoma: Setae nude; length of internal
humeral approximately 187 [i, slightly longer than first interspace;
external humeral, 206 \i in length; sacrals and clunals respectively.

A Revision of the Mite Family Bdellidae 399

in gently and strongly curving transverse rows. Anal region: Bor-
der striae parallel; anal setae and postanals wanting; 2 pairs of par-
anals. Genital region: Each genital plate wtih 6 equal, long setae
in linear arrangement; 3 pairs of paragenital setae; genital discs
small, caudal pair near termination of vestibule, distant from
anterior pairs; ovipositor with 20 setae of indeterminable position.
Legs (figs. 99, 100): Claws with 5 lateral rays each, row of minute
rays wanting; measurements: tibia I, 119 \j.; tarsus I, 349 [/.; tibia
II, 129 [i; tarsus II, 349 [>.. Chaetotaxy: coxae I-IV, 3, 2, 3, 2 tactile
setae; trochanters I-IV, 1 tactile seta each; basifemora I-IV, 12, 11,
9, 2 tactile setae; telofemora I-IV, 5, 5, 4, 4 tactile setae; genu I,
4 tactiles, 3 attenuate sensory setae; genu II, 4 tactiles, 1 attenuate
sensory seta; genu III, 4 tactiles, 1 attenuate sensory seta; genu IV,
4 tactiles; tibia I, 8 tactiles, 3 attenuate sensory setae, trichoboth;
tibia II, 8 tactiles, 1 attenuate and 1 blunt sensory seta, trichoboth;
tibia III, 8 tactiles, 1 attenuate sensory seta; tibia IV, 12 tactiles,
trichoboth; tarsus I, 20 ventral setae arranged in 2 rows, distal
15 plumose, 10 lateral and 2 dorsal tactile setae, 1 attenuate and 2
blunt sensory setae, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus
II, 18 ventrals, 10 laterals, 1 dorsal tactile seta, 2 blunt sensory
setae, 1 peg, dorsoterminals as in tarsus I; tarsus III, 17 ventrals,
9 laterals, trichoboth, 1 long, attenuate peg distal to trichoboth;
dorsoterminals as in tarsus I; tarsus IV, 20 xentrals, 2 lateral and
4 long, dorsolateral tactile setae, trichoboth, dt 1 reduced to one
solid, nude seta, dt 2, 3 solid, plumose.

Male: Unknown.

Holotype: Female, 2 miles west of Oakville, Napa Co., California,
December 31, 1953, G. A. Marsh, V. D. Roth, R. O. Schuster,
"Laurel association."

Location of type: The Snow Entomological Museum.

Remarks: The posterior sensilla are reduced as in other Thori-
hdella species, but lateral propodosomal setae are present. In known
species of the Odontoscirinae, only Octohdcllodes species and tin's
new Thoribdella species are characterized as having the lateral
propodosomal setae present. Due to the unusual presence of these
setae, this new species is called insolita. Drawings of the holotype.

Thoribdella communis, sp. nov.
(Figs. 22, 48, 97, 98, 146)
This species appears to be related to T. truncafa, sp. nov., but the
internal humerals are as long as the first interspaces and the palpal

400 The University Science Bulletin

basifemur is longer than the tibiotarsus. In T. truncota the internal
humerals are about one third of the first interspaces and the palpal
basifemur is three fourths the length of the tibiotarsus.

Female: Color in life dark red. Body robust, ovoid, weakly
constricted; length, including gnathosoma, 1730 [jl. Gnathosoma:
Length, 775 [x; palpus ( fig. 48 ) with basifemur longer than tibiotar-
sus; measurements: I, 27 pi; II, 265 [jl; III, 44 [jl; IV, 51 [x; V,
221 pi; des, 267 [x; ves, 197 [l. Chelicera (fig. 22) normal, nonstri-
ated, 447 [x in length; movable digit with inner surface apically
flattened, fixed digit with small subapical tooth, setae as figured.
Gnathosomal base striated, buccal cone nonstriated, punctate; ven-
tral setae forming two longitudinal series, vh 1-5 equidistant on
proximal one third of cone, vh 6 premedial between apex and vh
5; dorsal hypostomal setae approximately equal in length to vh 3.
Dorsal propodosoma: Striae finely broken; pattern similar to fig.
142; posterior sensilla approximate to base of median propodo-
somals, 36 [x in length; enlarged bases of median propodosomals
pilose (fig. 146); length of median propodosomals approximately
364 [X; eyes separated by a distance equal to 3 diameters of anterior
pair; distance between anterior sensilla, 87 [x. Dorsal hysterosoma:
Setae nude; length of internal humeral, 170 [x, approximately three
fourths of first interspace; external humeral about 175 [x in length;
sacrals in gently curving transverse row, external clunals absent.
Anal region: Border striae parallel; 2 pairs of anals, 1 pair of paran-
als; postanals wanting. Genital region: Each genital plate with
7 equal genital setae in linear arrangement; three pairs of para-
genitals; genital discs small, equidistant, in central area of vesti-
bule walls; ovipositor with 16 subapical, 4 postmedial setae. Legs
(figs. 97, 98): Claws with 6-7 lateral rays each, row of minute
rays wanting; measurements: tibia I, 136 \i; tarsus I, 359 [x; tibia

11, 138 [x; tarsus II, 369 \).. Chaetotaxy: coxae I-IV, 4, 3, 4, 2 tactile
setae; trochanters I-IV, 1 tactile seta each; basifemora I-IV, 13,

12, 8, 4 tactile setae; telofemora I-IV, 8, 8, 7, 6 tactile setae; genu
I, 6 tactiles, 5 attenuate sensory setae; genu II, 5 tactiles, 4 at-
tenuate sensory setae; genu III, 5 tactiles, 2 attenuate sensory setae;
genu IV, 5 tactiles, 2 attenuate sensory setae; tibia I, 10 tactiles,
3 attenuate sensory setae, 1 peg, trichoboth; tibia II, 10 tactiles,
1 attenuate and 1 blunt sensory seta, trichoboth; tibia III, 11 tactiles,
1 attenuate sensory seta; tibia IV, 12 tactiles, trichoboth; tarsus
I, 22 plumose ventrals arranged in 2 rows, 10 lateral and 2 dorsal
tactile setae, 2 attenuate and 2 blunt sensory setae, 1 peg, dt 1 solid.

A Revision of the Mite Family Bdellidae 401

nude, dt 2, 3 solid, plumose; tarsus II, 21 ventrals as in tarsus I,
10 lateral tactiles, 1 dorsal tactile, 1 attenuate and 2 blunt sensory
setae, 1 peg, dorsoterminals as in tarsus I; tarsi III-IV, 21 ventrals as
in tarsus I, 8 laterals, trichoboth, dt 1 reduced to 1 solid, nude seta,
dt 2, 3 solid, plumose on tarsus IV.

Male: Identical to female except larger; length, including gnatho-
soma, 1560 p.; amphioid sclerites with 3, 4, 2 setae each.

Holotype: Male, west slope of Cortez Pass, Mexico, Mexico,
August 11, 1954, W. T. Atyeo, under rock, altitude 13,000 feet.

Allotype: Female, west slope of Cortez Pass, Mexico, Mexico,
August 11, 1954, W. T. Atyeo, moss (Berlese funnel extraction).

Paratypes: Two males, four females, same data as holotype; five
males, twelve females, 4 miles west of San Cristobal de las Casas,
Mexico, Mexico, July 11, 1955, R. E. Beer, under rocks; two males,
5 miles south of Avila Camacho, Distrito Federal, Mexico, July 3,
1956, R. E. Beer, on bunch grass, Epicamees sp.; one male, 16
miles north of Juchitan, Oaxaca, Mexico, July 4, 1955, R. E. Beer,
under rock; one female, 7 miles north of Cuernavaca, Morelos,
Meixco, August 14, 1954, W. T. Atyeo, beating lichens, altitude
9,000 ft.; one female, twenty miles west of Mexico (city), Mexico,
July 31, 1954, W. T. Atyeo, under rock, altitude 9,000 ft.; one female,
7 miles northeast of Jacala, Hidalgo, Mexico, July 27, 1954, W. T.
Atyeo, under log.

Location of types: The holotype, allotype, and twenty-three para-
types deposited in the Snow Entomological Museum. Two para-
types deposited in each of the following: the South Australian
Museum, the British Museum (Natural History), and the United
States National Museum.

Remarks: The total length of the majority of the specimens is
between 1350 [x to 1700 pi, however, one female from the San Cris-
tobal series measures about 2200 [jl. Other measurements of the
larger specimens are proportional to total length, for example, the
palpal measurements of the extremely large female are: I, 30 [x; II,
337 [l; III, 63 ix; IV, 66 [i; V, 267 ix; des, 315 [x; ves, 255 |x.

The number of setae on the palpal basifemur varies from six to
nine, and on the tibiotarsus, two specimens of the paratype series
have a fourth seta in the median proximal row. Variations in the
number of tactile setae on the leg segments are slight, and there
may be five or six attenuate sensory setae on genu I. Individuals
of this species are abundant in Mexico, hence the name of coin-
7nunis. Drawings of the holotype.

402 The University Science Bulletin

Thoribdella truncato, sp. nov.

(Figs. 24, 50, 101, 102, 149, 150)
This species is related to T. communis, sp. nov., but has highly
modified posterior sensilla (fig. 150) and internal humerals one
third the length of the first interspaces, rather than small, setaceous
sensilla and internal humerals equal to the first interspaces.

Female: Color in life unknown; body ovoid, weakly con-
stricted; length, including gnathosoma, 1560 ^.. Gnathosoma:
Length, 384 [x; palpus (fig. 50) with basifemur shorter than tibio-
tarsus; measurements: I, 17 [x; II, 153 [x; III, 53 [x; IV, 53 \).; V,
182 [i; des, 122 [x; ves 97 [jl. Chelicera (fig. 24) normal, nonstriated,
333 pi. in length; movable digit with inner surface distally flattened;
setae as figured. Gnathosomal base and proximal third of buccal
cone striated; distal two thirds of cone nonstriated; v>h 1-6 in two
longitudinal series, vh 1 inserted between palpal articulations, vh 1-5
equidistant, vh 6 approximately median between vh 5 and apex;
dorsal hypostomal setae half the length of vh 1. Dorsal propodo-
soma: Striae finely broken; pattern similar to fig. 142; median
propodosomals (fig. 149), 289 \). in length; posterior sensilla, 12 \i in
length, modified as in fig. 150; eyes separated by distance equal to
two diameters of anterior pair; distance between anterior sensilla,
83 [A. Dorsal hijsterosoma: Setae finely pilose; length of internal
humeral, 65 \i, approximately one third of first interspace; external
humeral, 65 [j. in length; sacrals and clunals respectively in gently
and strongly curving transverse rows. Anal region: Border striae
parallel; one pair of anal setae; paranals wanting; postanals flank-
ing termination of cleft, shorter than clunal setae. Genital region:
Eight small, unequal setae in linear arrangement, anterior 3 longer
than posterior 5, two pairs of paragenitals; genital discs small, equi-
distant; ovipositor with 16 subapical, 4 postmedial setae. Legs ( figs.
101, 102 ) : Claws with 6-7 lateral rays, one row of minute rays each;
measurements: tibia I, 121 [i.; tarsus I, 289 \i; tibia II, 122 [x; tarsus

II, 291 a. Chaetotaxy: coxae I-IV, 4, 4, 5, 4 tactile setae; trochanters
I-IV, 1 tactile seta each; basifemora I-IV, 17, 13, 10, 6 tactile setae;
telofemora I-IV, 8, 7, 6, 5 tactile setae; genu I, 5 tactiles, 7 attenuate
sensory setae; genu II, 6 tactiles, 6 attenuate sensory setae; genu

III, 5 tactiles, 5 attenuate sensory setae; genu IV, 6 tactiles, 3 at-
tenuate sensory setae; tibia I, 12 tactiles, 5 attenuate sensory setae,
1 blunt sensory seta, 1 apically forked peg, trichoboth; tibia II,

A Revision of the Mite Family Bdellidae 403

12 tactiles, 3 attenuate sensory setae, 1 blunt sensory seta, tricho-
both; tibia III, 13 tactiles, 6 attenuate sensory setae; tibia IV, 14
tactiles, trichoboth; tarsus I, 36 plumose ventrals arranged in 2
rows proximally, to five irregular series distally, 15 lateral and 2
dorsal tactiles, 2 attenuate and 2 blunt sensory setae, 1 peg, dt 1,
solid, nude, dt 2, 3 solid, plumose; tarsus II, 36 ventrals as in tarsus
I, 12 lateral and 2 dorsal tactiles, 1 attenuate and 2 blunt sensory
setae, 1 peg, dorsoterminals as in tarsus I; tarsus III, ventrals,
laterals, dorsoterminals as in tarsus I, trichoboth; tarsus IV, ven-
trals, laterals, dt 2, S as in tarsus I, trichoboth, dt 1 reduced to 1
solid, nude seta.

Male: Identical to female except in general size; total length,
including gnathosoma, 1350 \j., other measurements vary accord-
ingly; amphioid sclerites with 3, 1, 3, 2 setae each.

Holotijpe: Male, 38 miles northeast of Montemorelos, Nuevo
Leon, Mexico, August 4, 1955, R. E. Beer, under rock.

Allotype: Female, same data as holotype.

Paratypes: One male, same data as holotype; two females, 3 miles
west of Antiguo Morelos, Tamaulipas, Mexico, R. E. Beer, August
3, 1955, under rocks; one male, Teziutlan, Pueblo, Mexico, July 22,
1955, R. E. Beer, hand picked from litter on forest floor; one female.
Vera Blanca, Costa Rica, entre los volcanes Poas y Barba, altitude
2,000 m. s. m., August, 1938, Alex Bierig.

Location of types: The Snow Entomological Museum.

Remarks: In the Mexican series, the only notable variation is
the number of attenuate sensory setae on tibia II. The specimens
from Montemorelos and one from Antiguo Morelos have three at-
tenuate sensory setae on tibia II of one side of the body and tsvo
setae on the other side; in the other specimens, the number of setae
is the same on both sides, either two or three.

The Costa Rican specimen is larger than the Mexican specimens,
measuring 1880 \i in total length. This specimen has a simple peg
and nine attenuate sensory setae on tibia I, rather than a bifurcate
peg and five attenuate sensory setae. Other segments of the legs
also have an increased number of setae, while the genital plates have
nine pairs of setae, instead of eight.

To indicate the unique modification of the posterior scnsilla,
this species is named truncata. Drawings of the holotype.

404 The University Science Bulletin

Thoribdella meridionalis (Thor)

(Figs. 21, 51, 95, 96, 145)

Biscirus (Biscirus) meridionalis Thor, 1931, Zool. Anz., vol. 92, no. Yz, pp. 74-76.

This species is closely related to Thoribdella norvegicus (Thor),
1913 ( = Biscirus ( B. ) norvegicus), but is much smaller, as indi-
cated by the palpal measurements given for norvegicus (Thor,
1931) : I, 25 [x; II, 460 \i; III, 88 [>.; IV, 136 ix; V, 33 [x. The relative
lengths of the palpal telofemur and genu may be used to diflFeren-
tiate these species, T. meridionalis having the segments approxi-
mately equal, and norvegicus with the fourth segment almost twice
as long as the third.

Female: Color unknown; length, including gnathosoma, 866 [x
(852-1143 [x). Gnathosoma: Length, 218 [x (218-277 ix); palpus
(fig. 51) short, about half of tibiotarsus extending beyond hypo-
stome; measurements: I, 12 [x (12-15 [x); II, 94 [x (94-129 jx);
III, 22 [X (20-26 [x); IV, 24 [X (24-31 [x); V, 78 [x (71-109 [x); des,
107 tx (107-129 [x); ves, 97 [x (94-117 [x). Chelicera (fig. 21) nor-
mal, nonstriated, 204 [x (201-247 (x) in length; movable digit flat-
tened distally on inner surface; setae as figured. Gnathosomal base
striated, buccal cone nonstriated; vh 1-5 equidistant, in 2 longi-
tudinal series on proximal third of cone, vh 6 mesad to series, mid-
way between vh 5 and apex of cone; dorsal hyi:)ostomal setae half
the length of vh 2. Dorsal propodosoma: Striae finely broken, pat-
tern similar to fig. 142; median propodosomals (fig. 145) enlarged
basally, 162 [x (158-184 [x) in length; posterior sensilla thin, ap-
proximate to medial propodosomals, 17 [x (15-17 jx) in length; eyes
separated by distance equal to two and one-half diameters of an-
terior pair; distance between anterior sensilla, 53 [x (44-53 [x).
Dorsal hijsterosoma: Setae nude; length of internal humeral, 63 [x
(51-68 ]}.), approximately one half of first interspace; external
humeral, 66 jx (53-68 [x) in length; sacrals and clunals respectively
in gently and strongly curving transverse rows. Anal region: Border
striae parallel, anals wanting, 1 pair of paranals, postanals flanking
termination of cleft, shorter than clunals. Genital region: Each
genital plate with 5 (5-7) small, equal setae in linear arrangement;
3 pairs of paragenitals; ovipositor with 16 subapical, 4 postmedial
setae, l^egs (fig. P6): Caws with 4-5 lateral rays each, row
of minute rays wanting; measurements: tibia I, 53 [x (53-65 (x);
tarsus I, 136 -x (136-179 jj.); tibia II, 53 [x (53-66 [x); tarsus II,
134 [X (134-170 [x). Chaetotaxy: coxae I-IV, 4, 2, 3, 2 tactile setae;
trochanters I-IV, 1 tacti'e seta each; basifemora I-TV, 9, 7, 5, 3 tac-

A Revision of the Mite Family Bdellidae 405

tile setae; telofemora I-IV, 5, 5, 4, 4 tactile setae; genu I, 4 tactiles,
5(6) attenuate sensory setae; genu II, 4 tactiles, 3 attenuate sensory
setae; genu III, 4 tactiles, 2 (3) attenuate sensory setae; genu IV,
4 tactiles, 3 attenuate sensory setae; tibia I, 7 tactiles, 3 attenuate
sensory setae, 1 peg, trichoboth; tibia II, 6 tactiles, 1 attenuate and
1 blunt sensory seta, trichoboth; tibia III, 7 tactiles, 1 attenuate
sensory seta; tibia IV, 7 tactiles, trichoboth; tarsus I, 12 plumose
ventrals in 2 rows, 6 lateral and 1 dorsal tactile setae, 2 attenuate
and 2 blunt sensory setae, 1 peg, dt 1 solid, nude, dt 2, 3 solid,
plumose; tarsus II, similar to tarsus I but lacking the 2 attenuate
sensory setae; tarsus III, 13 plumose ventrals, 5 laterals, trichoboth,
dorsoterminals as in tarsus I; tarsus IV, 15 ventrals, 4 laterals,
trichoboth, dt 1 reduced to 1 solid, nude seta, dt 2, 3 solid, plumose.
Male: Not examined.

Type: Male, near Tange, North Africa, January, 1931, F. Grand-
jean, vegetable debris under bush.

Location of type: Unknown.

Remarks: The redescription is based on thirteen females collected
in the United States ( Maryland, Connecticut, Kansas, Michigan,
California), Iceland, Sweden (Vasterbotten), and Germany (Ost-
Holstein). The specimens from Germany have an extra seta on
the palpal tibiotarsus, but other than this exception, all the speci-
mens coincide with the redescription. Illustrations were prepared
from a female with the following data: Patuxent Wildlife Refuge,
Maryland, R. O. Drummond, February 23, 1955, Peromyscus leii-
copus novcboracensis nest, no. 27.

Thvrihdella simplex, sp. nov.

(Figs. 8,23,49, 103, 104, 147)

The approximately equal lengths of the palpal telofemur and
genu distinguish this species from the related T. californica, in
which the genu is twice as long as the telofemur.

Female: Color unknown. Body ovoid, weakly constricted;
length, including gnathosoma, 2100 pi. Gnatlwsoma: Length, 572 jjl;
palpus (fig. 49) with tibiotarsus extending beyond hypostome;
measurements: I, 31 [x; II, 289 ^- III, 68 ^■, IV, 88 |x; V, 292 jx; dcs,
218 [i; ves, 167 [a. Chelicera (figs. 8, 23) normal, nonstriated, 568 (x
in length; inner surface of movable digit flattened distally; setae as
figured. Gnathosomal base and buccal cone to level of vh 3 stri-
ated, distal portion of cone nonstriated; vh 1-6 in 2 longitudinal
series, vh 1, 2 between palpal articulations, vh 1-4 equidistant, vh

406 The University Science Bulletin

5 slightly removed distally, vh 6 medial between vh 5 and apex;
dorsal hypostomal setae longer than any member of vh series.
Dorsal propodosoma: Striae finely broken, pattern similar to fig.
142; median propodosomal basally expanded (fig. 147), 349 [t. in
length; posterior sensilla thin, 36 \i in length, approximate to median
propodosomals; eyes separated by distance equal to two diameters
of anterior pair; distance between anterior sensilla, 104 pi. Dorsal
hysterosoma: Setae plumose; length of internal humeral, 85 [j,, ap-
proximately one third of first interspace; external humeral, 94 [a
in length; sacrals and clunals respectively in gently and strongly
curving transverse rows. Anal region: Border striae joarallel; anal
setae wanting; one pair of paranals; postanals flanking termina-
tion of cleft, shorter than clunals. Genital region: Each genital
plate with 9 unequal setae in linear arrangement, anterior 3
slightly larger than posterior 6 setae; 3 pairs of paragenitals; gen-
ital discs equidistant, anterior 2 pairs large, posterior pair small;
ovipositor with 16 subapical, 4 postmedial setae. Legs (figs. 103,
104): Claws with 6-7 lateral rays each, row of minute rays want-
ing; measurements: tibia I, 218 [j.; tarsus I, 394 [jl; tibia II, 227 pi;
tarsus II, 404 ^. Chaetotaxy: coxae I-IV, 4, 3, 4, 3 tactile setae;
trochanters I-IV, 1 tactile seta each; basifemora I-IV, 16, 15, 12, 5
tactile setae; telofemora I-IV, 11, 10, 10, 6 tactile setae; genu I, 7
tactiles, 12 attenuate sensory setae; genu II, 6 tactiles, 6 attenuate
sensory setae; genua III-IV, 6 tactiles, 4 attenuate sensory setae
each; tibia I, 13 tactiles, 14 attenuate sensory setae, 1 blunt sensory
seta, 1 peg, trichoboth; tibia II, 14 tactiles, 6 attenuate sensory
setae, 1 blunt sensory seta, trichoboth; tibia III, 13 tactiles, 1 atten-
uate sensory seta; tibia IV, 14 tactiles, trichoboth; tarsus I, 48 plu-
mose ventrals arranged in 4 irregular rows, 12 lateral and 2 dorsal
tactile setae, 2 attenuate and 2 blunt sensory setae, 1 peg, dt 1 solid,
nude, dt 2, 3 solid, plumose; tarsus II, similar to tarsus I, except
lacking 1 lateral tactile seta and 2 attenuate sensory setae; tarsus
III, 44 ventrals, 10 laterals, trichoboth, dorsoterminals as in tarsus
I; tarsus IV, 44 ventrals, 9 laterals, trichoboth, dt 1 reduced to 1
solid, nude seta, dt 2, 3 solid, plumose.

Male: Identical to female except in the chaetotaxy of the palpal
basifemur, the number of attenuate sensory setae on legs I and II,
and general size. Measurements: total length, 1605 [).; palpus:
I, 25 [i; II, 224 \l; III, 43 [x; IV, 56 [;.; V, 272 -x; des, 213 jj.; ves, 158
[J.; tibia I, 143 [>.; tarsus I, 313 p.; tibia II, 146 [x; tarsus II, 322 [;.; che-
licera, 412 p.; gnathosoma, 433 \i.; median propodosomals, 328 ij.; pos-
terior sensilla, 34 it.; distance between anterior sensilla, 77 [j.. Palpal

A Revision of the Mite Family Bdellidae 407

basifemur with 5 setae; each genital plate with 10 setae; amphioid
sclerites with 3, 1, 3, 1 setae each; genua I-II, 15 tactile setae, 9 at-
tenuate sensory setae; tibiae I-II, 15 tactiles, 7 attenuate sensory
setae. Tactile setae of legs slightly reduced from numbers indicated
in holotype description.

Holotype: Female, Clinton, Douglas Co., Kansas, May 5, 1955,
D. S. Narayan, grass (Berlese funnel extraction).

Allotype: Male, 1 mile west of Monticello, Madison Co., Flor-
ida, April 7, 1957, W. T. Atyeo, leaf litter.

Paratypes: One female, same data as holotype; four females,
same data as allotype; two females, O'Leno State Park, Florida,
April 6, 1957, W. T. Atyeo, leaf litter; one male, 2 females col-
lected at Dinuba, Tulare Co., California, by Francis M. Summers
as follows: one male, March 12, 1945, garden soil; one female,
March 17, 1945, mulch; one female, March 26, 1945, mulch.

Location of types: Holotype, allotype, and seven paratypes de-
posited in the Snow Entomological Museum; one paratype at each
of the following: the United States National Museum, the South
Australian Museum, and the British Museum (Natural History).

Remarks: The Florida and California paratypes are approxi-
mately 1800 PL in length, slightly smaller than the holotype. In each
of these specimens, there is a reduction in the number of setae; the
palpal tibiotarsus has eight to nine setae, the palpal basifemur has
five to six setae, and the genital plates have seven to eight setae
each. The tactile setae on the leg segments (except the trochanters)
and the attenuate sensory setae on the genua and the tibiae of
legs I and II vary slightly from the numbers indicated in the de-
scription of the holotype. This species is named simplex because
of the lack of striking morphological characters. Drawings of leg
II, palpus, and dorsal propodosoma of the holotype; leg I drawn
frm the Kansas paratype.

Genus Octohdellodes, gen. nov.

Type: Octohdellodes htirdi sp. nov.

This new genus is closely related to Bdellodes, but has the lateral
propodosomal setae present and six or seven pairs of ventral hypos-
tomal setae. The chelicerae are normal and each has two setae.
The movable digits are smooth and longer than the fixed digits.
The posterior pseudostigmatic organs are unmodified and the pos-
terior sensilla are longer than the median propodosomal setae.
Legs II are shorter than legs I.

408 The University Science Bulletin

Key to Species of Octobdellodes

1. Palpal genu with four setae; palpal genu and telofemur approxi-
mately equal in length hurdi p. 408

Palpal genu with seven setae; palpal genu twice as long as the
telofemur infrequens p. 410

Octobdellodes hurdi, sp. nov.
(Figs. 18, 45, 93, 94, 152)

This species is related to Octobdellodes infrequens, sp. nov., but
has the palpal genu and telofemur approximately equal in length,
rather than the genu twice as long as the telofemur.

Female: Color in life unknown. Body narrowly ovoid, weakly
constricted; length including gnathosoma, 1380 ^.. Gnathosoma:
Length, 374 \i; palpus (fig. 45) with tibiotarsus extending beyond
hypostome; measurements: I, 17 [jl; II, 189 p.; Ill, 32 \i; IV, 34 ;/.;
V, 195 pi; des, 133 [j.; ves 128 [i. Chelicera (fig. 18) normal, non-
striated, 323 \i in length; chela smooth, fixed digit shorter than
movable digit; setae as figured. Gnathosomal base striated, buccal
cone nonstriated; vh 2-6 in longitudinal series, vh 1, 2 form trans-
verse row on striated base, vh S between palpal articulations, vh
3-5 equidistant, vh 6 median between apices of lateral lips and vh
5; dorsal hypostomal setae equal in length to vh 2. Dorsal propodo-
soma ( fig. 152 ) : Striae finely broken; lateral propodosomals nude,
70 [i in length; median propodosomals nude, 85 [x in length; eyes
subequal, interval between eyes equal to three diameters of the
larger, anterior pair; distance between anterior sensilla, 68 [x. Dor-
sal htjsterosoma: Setae nude; length of internal humeral, 68 \>.,
approximately one third of first interspace; external humeral, 73 \i
in length; sacrals in gently curving transverse row, clunals in sub-
rectangular arrangement. Anal region: Border striae parallel;
anal setae wanting; two pairs of paranals; postanals wanting. Geni-
tal region: Each genital plate with 7 small, equal setae in linear
arrangement; 4 pairs of paragenitals; genital discs small, ovi-
positor with 14 subapical, 6 postmedial setae. Legs (figs. 93, 94):
Claws with 4 lateral rays each, minute rays wanting; measurements:
tibia I, 94 {x; tarsus I, 188 \i; tibia II, 80 [x; tarsus II, 172 [x. Chaeto-
taxy: coxae I-IV, 6 tactile setae each; trochanters I-IV, 1, 2, 2, 2
tactile setae; basifemora I-IV, 9, 9, 6, 3 tactile setae; telofemora
I-IV, 7, 7, 6, 5 tactiles; genu I, 5 tactiles, 5 attenuate sensory setae;
genua II-IV, 3 tactiles, 2 attenuate sensory setae; tibia I, 7 tactiles.

A Revision of the Mite Family Bdellidae 409

3 attenuate sensory setae, 1 peg, trichoboth; tibia II, 7 tactiles, 1
attenuate and 1 blunt sensory seta, trichoboth; libia III, 8 tactiles,

1 attenuate sensory seta; tibia IV, 8 tactiles, trichoboth; tarsus I,
15 pkimose ventrals in two rows, 7 lateral and 2 dorsal tactile setae,

2 attenuate and 2 blunt sensory setae, 1 capitate peg, dt 1 Eolid,
nude, dt 2, 3 solid, plumose; tarsus II, 15 plumose ventrals, 5 lateral
and 1 dorsal seta, 1 attenuate and 2 blunt sensory setae, 1 peg,
dorsoterminals as in tarsus I; tarsus III, 15 plumose ventrals, 6
laterals, trichoboth, dorsoterminals as in tarsus I; tarsus IV, 15
plumose ventrals, 5 laterals, trichoboth, dt 1 reduced to 1 solid,
nude seta, dt 2, 3 solid, plumose.

Male: Identical to female except in genital region; each genital
plate with 8 setae; amphioid sclerites with 3, 4, 1 setae each.

Holotijpe: Female, Point Barrow, Alaska, July 5, 1952, P. D. Hurd,
Berlese intermediate.

Allotype: Male, same data as holotype.

Paratypes: Four males and nine females collected by P. D. Hurd,
at Point Barrow, Alaska, in 1952; these include: three females with
same data as holotype; two males, four females, June 26, dry
timdra. above frost line, top of polygon (Berlese funnel extraction);
two males, one female, June 26, ring sample, frost scar on ridge; one
female, July 26, ridge plot.

Location of types: The holotype, allotype, and five paratypes
are deposited in the Snow Entomological Museum. Two paratypes
are deposited at the following: the South Australian Museum, the
British Museum (Natural History), the United States National
Museum, and the University of California.

Remarks: Total size is relatively uniform, the greatest variation
occurs in the numbers of setae. The holotype has six tactile setae
on the left coxa IV and nine setae On the right coxa. Genital setae
in the males vary from eight to nine pairs, and in the females, from
seven to eight pairs. Tactile setae on the leg segments vary from
two more, to two less, than the numbers indicated in the description
of the holotype. The proximal seta on the medial surface of the
palpal basifemur is absent in a few specimens. The species is
named for Dr. P. D. Hurd of the University of California, who
collected the type series of this species. Drawings are of the holo-
type.

410 The University Science Bulletin

Octobdellodes infrequens, sp. nov.
(Figs. 19, 20, 46, 89, 90, 153)

This species is related to O. luirdi, sp. nov., but can be distin-
guished by the palpal genu which is two times longer than the telo-
femur and which has seven setae rather than the genu equal in
length to the telofemur and with four setae.

Female: Color in life unknown. Body large, ovoid, weakly con-
stricted; length, including gnathosoma, 2700 \i. Gnathosoma:
Length, 629 [x; palpus (fig. 46) long, half of the genu and tibiotarsus
extending beyond hypostome; measurements: I, 30 [x; II, 357 \^; III,
68 [x; IV, 156 [x; V, 335 ^■, des, 323 pi; ves, 264 \i. Chelicera normal,
nonstriated, 575 [x in length; fixed digit and movable digit with pre-
apical tooth, fixed digit slightly shorter than opposing digit; chelic-
eral setae inserted as in figs. 19, 20. Gnathosomal base striated,
buccal cone nonstriated; vh 1, 2 form transverse row on hypostome
base, vh 2-5, 7 form longitudinal series, vh 6 mesad, vh 1 laterad;
vh 2-7 approximately equidistant; dorsal hypostomal setae about
half the length of vh 1. Dorsal propodosoma (fig. 153): Striae
finely broken; lateral propodosomals nude, 158 ;j. in length; median
propodosomals nude, 145 [x in length; eyes separated by distance
equal to 2 diameters of anterior pair; distance between anterior
sensilla, 68 [x. Dorsal hysterosoma: Setae nude; length of internal
humeral, 146 |x, approximately half of first interspace; external hu-
meral, 155 [X in length; sacrals in gently curving transverse row,
clunals in subrectangular arrangement. Anal region: Border striae
parallel; anal setae wanting; 2 pairs paranals; postanals removed
laterad of termination of cleft, shorter than clunals. Genital region:
Each genital plate with 9 long, equal setae in linear arrangement,
5 pairs of paragenitals; genital discs equidistant, in middle por-
tion of vestibule; ovipositor with 16 subapical, 6 postmedical setae.
Legs ( figs. 89, 90 ) : Claws with 6-7 lateral rays, 1 row of minute
rays each; measurements: tibia I, 187 [x; tarsus I, 377 jx; tibia II,
187 [x; tarsus II, 377 [x. Chaetotaxy: coxae I-IV, 5, 5, 6, 3 tactile setae:
trochanters I-IV, 1, 2, 2, 2 tactile setae; basifemora I-IV, 14, 15, 10,
5 tactile setae; telofemora I-IV, 10, 9, 9, 7 tactile setae; genu I, 6
tactile setae, 5 attenuate sensory setae; genu II, 6 tactile setae, 4
attenuate sensory setae; genua III-IV, 6 tactile setae and 3 attenu-
ate sensory setae each; tibia I, 11 tactiles, 4 attenuate sensory setae,
1 peg, trichoboth; tibia II, 11 tactiles, 3 attenuate and 1 blunt
sensory seta, trichoboth; tibia III, 14 tactiles, 1 attenuate sensory
seta; tibia IV, 13 tactiles, trichoboth; tarsus I, 32 plumose ventral

A Revision of the Mite Family Bdellidae 411

setae arranged in 2 rows on proximal third, in four rows on distal
two thirds, 10 lateral and 2 dorsal tactiles, 2 attenuate and 2 blunt
sensory setae, 1 peg, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus
II, 32 plumose ventrals arranged as in tarsus I, 10 lateral and 2
dorsal tactiles, 1 attenuate and 2 blunt sensory setae, 1 peg, dorso-
terminals as in tarsus I; tarsus III, ventrals, laterals, dorsoterminals
as in tarsus II, trichoboth; tarsus IV, 36 ventrals, 8 laterals arranged
as in tarsus I, trichoboth, dt 1 reduced to 1 solid, nude seta, dt 2,
3 solid, plumose.

Male: Identical to female except in size and chaetotaxy of the
palpus. Measurements: length, including gnathosoma, 1740 [i; pal-
pus: I, 26 ii; II, 325 [x; III, 54 [/.; IV, 104 [l; V, 269 [x; des, 247 [/.; ves,
207 ii; chehcera, 497 \i.; gnathosoma, 476 ix; tibia I, 143 [i; tarsus I,
320 [x; tibia II, 143 pi; tarsus II, 317 ix. Setae on the dorsal propo-
dosoma and dorsal hysterosoma only slightly shorter. Second
palpal segment with 10 tactile setae, fifth segment with 12 tactile
setae (excluding apical setae). Amphioid sclerites with 3, 4, 2
setae each.

Holotijpe: Male, Douglas County, Kansas, February 17, 1952,
C. C. Hall, on shagbark hickory.

AUotijpe: Female, Sioux City, Iowa, October 15, 1921, C. N.
Ainslie, under stone.

Paratypes: Male, Imoden, Arkansas, received at USNM Janu-
ary 14, 1935, from B. C. Marshall; female, Patuxent Wildlife
Refuge, Maryland, January 13, 1954, R. O. Drummond, Peromyscus
leucopus novehoracensis nest, no. 11.

Location of types: The holotype and paratypes are deposited in
the Snow Entomological Museum; the allotype is deposited at the
United States National Museum.

Remarks: It is difficult to estimate the amount of variation in the
species because of the limited number of specimens. The allotype
is an extremely large female, probably gravid, while the paratypes
are comparable in size to the holotype. The number of ventral
hypostomal setae in the four specimens is very variable. Five pairs
of setae are always present on the buccal cone, but in the transverse
row across the gnathosomal base, there may be two to three pairs,
totaling therefore, seven to eight pairs of ventral hypostomal setae
rather than the usual six pairs.

This species, although apparently widely distributed, has been
infrequently collected, therefore the sj^ecific name of infrcqtiens
has been selected for this new taxon. Drawings of the holotype.

412 The University Science Bulletin

Genus Bdellodes Oiidemans

Bdellodcs Oudcmans, 1937 ( = Srirw.s, sensit Thor, 1931, non Herniann, 1804),
Kritisch Historisch Overzicht der Acarologie, Leiden, vol. 3, part C, p. 1217.
(Type: Scirus longirostris Ilemiann, 1804 [by original designation].)

Hoplosciru.'! Thor, 1937, Zool. Anz., vol. 119, no. %, p. 43 (new synonym).
(Type: Scirus dnhitatus Womersley, 1933 [by original designation].)

Bdellodes is closely related to Octohdellodes, gen. nov., but lacks
the lateral propodosomal setae. The chelicerae are normal and
each has one or two setae. The sickle-shaped movable digits are
longer than the fixed digits, and may be smooth, flattened distally
on the inner surfaces, or have a single tooth. The posterior pseu-
dostigmatic organs are unmodified and the posterior sensilla are
longer than the median propodosomal setae. Legs II are usually
shorter than legs I.

The genus Hoploscirus was erected for species of Bdellodes
(= Scirus, sensu Thor) with distinct, reniform dorsal shields. The
differentiating character for this group is an extreme modification
of cuticular deposition, which is considered to be a continuous
character.

Key to the Species of Bdellodes

1. Chelicera with one seta; palpal basifemur and tibiotarsus (ex-
cluding end setae) with twelve to fourteen setae each longirostris p. 412
Chelicera with two setae; palpal basifemur and tibiotarsus (ex-
cluding end setae) with five to six setae each hisetosa p. 414

Bdellodes longirostris (Hermann)
(Figs. 14, 42, 87, 88)

Scirus longirostris Hermann, 1804, Mem. Apt. p. 62.

Bdella peregrina Banks var. iowaensis Ewing, 1917, Bull. Amer. Mus. Nat.
Hist., vol. 37, p. 150 (new synonym).

This species is closely related to Bdellodes porrecftis (Kramer'^
1898, but the palpal genu and telofemur are approximately equal
in length and each digit of the chela has one preapical tooth, rather
than the genu twice as long as the telofemur and the digits smooth.

Female: Color in life red, with dark blue flecks. Body ovoid;
length, including gnathosoma, 1810 \i ( 1335-2270 \i ) . Gnathosoma:
Length, 504 |i, (447-568 [x); palpus (fig. 42) with short, subequal
end setae; measurements: I, 27 [jl (24-28 pi); II, 357 [jl (264-408 n);
III, 61 [x (60-63 [i); IV, 70 IX (50-70 jx); V, 325 [x (24.3-352 [x); des,
218 tx (197-238 [x); ves, 201 [x (179-201 [;.). Chelicera (fig. 14) nor-
mal, nonstriated, 471 \x (412-547 ix) in length; each digit with one
preapical tooth; single seta inserted at approximately two thirds
of the length of chelicera from base. Gnathosomal base striated,

A Remsion of the Mite Family Bdellidae 413

buccal cone nonstriated; vh 2-6 form 2 longitudinal series, vh 1
laterad, between articulations of palpi; vh 2-5 approximately equi-
distant, vh 6 median between vh 5 and termination of buccal cone;
dorsal hypostomal setae approximately equal in length to vh 3.
Dorsal propodosoma: Striae finely broken, pattern similar to fig.
153; median propodosomals finely plumose, 133 [). (119-163 \i.) in
length; eyes separated by distance equal to 2 diameters of anterior
pair, interval between eyes with transverse striae; distance between
anterior sensilla, 85 [x (65-88 pi). Dorsal hijsterosoma: Setae finely
plumose; length of internal humeral, 99 [a ( 87-109 [x ) , approximately
half of first interspace; external humeral, 100 \i (95-128 [>.) in length;
sacrals in gently curving transverse row, clunals in subrectangular
arrangement. Anal region: Border striae parallel; 1 pair of anal
setae; paranals wanting; postanals flanking cleft approximately one
fourth length of cleft from dorsal termination, shorter than clunals.
Genital region: Each plate with 6(7) long, equal, attenuate setae
in linear arrangement; 4(5) pairs of paragenitals, anterior pair at
level of anterior termination of aperture; genital discs small, one
pair at anterior end, two pairs approximate, caudad; ovipositor
with 18 subapical setae. Legs (figs. 87, 88) : Claws with 6-7 lateral
rays and one row of minute rays each; measurements: tibia I, 167 \i.
(126-177 [x); tarsus I, 296 \i ( 264-325 [x ) ; tibia II, 167 [x ( 126-175 ix);
tarsus II, 296 [). (264-322 -x). Chaetotaxy: coxae I-IV, 5 (4), 3, 4, 2
tactile setae; trochanters I-IV, 1 tactile seta each; basifemora I-IV,
15 (13-16), 15 (14), 13 (9-13), 6 (4-6) tactile setae; telofemora
I-IV, 8 (7), 7, 6, 6 (7) tactile setae; genu I, 6 (5) tactile setae, 5
(4-6) attenuate sensory setae; genu II, 6 tactile setae, 3 (2) at-
tenuate sensory setae; genua III-IV, 6 (5) tactile setae and 2 at-
tenuate sensory setae each; tibia I, 13 (10-14) tactile setae, 4 (3)
attenuate sensory setae, 1 peg, trichoboth; tibia II, 12 (9-12) tactile
setae, 1 attenuate and 1 blunt sensory seta, trichoboth; tibia III,
12 (11) tactile setae, 1 attenuate sensory seta; tibia IV, 13 (10-13)
tactile setae, trichoboth; tarsus I, 29 (24-32) plumose ventral setae,
distal members in 4 irregular rows, proximal members in 2 rows,
11 (10) lateral and 2 dorsal tactile setae, 2 attenuate and 2 bhint
sensory setae, 1 peg, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus
II, similar to tarsus I except with 9 (7-9) laterals, 1 dorsal tactile
seta, 1 attenuate sensory seta; tarsus III, ventrals and dorsoterminals
as in tarsus I, 8 laterals, trichoboth; tarsus IV, ventrals as in tarsus
I, 9 laterals, trichoboth, dt 1 reduced to one solid, nude seta, dt 2, 3
solid, plumose.

414 The UNivERSiTi' Science Bulletin

Male: Identical to female; amphioid sclerites with 3, 4, 2 setae
each.

Type: Europe.

Location of type: Unknown.

Remarks: Specimens examined were collected in: Mexico (Mi-
choacan, Jahsco, Oaxaca, Guanajuato, Mexico, Distrito Federal,
Puebla, Nuevo Leon, Guerrero, Tamaulipas), the United States
(Texas, California, Kansas, Arkansas, Florida, Michigan, Montana),
Cuba, Costa Rica, Jamaica, Argentina, and Denmark. The number
of setae on the palpus may vary from ten to fifteen on the basifemur,
and from twelve to fourteen on the tibiotarsus. Redescription and
drawings are based on two males from: twenty miles west of
Morelia, Michoacan, Mexico, July 20, 1956, R. E. Beer, under rock.

Bdellodes hisetosa, sp. nov.

(Figs. 47,91,92, 155)

This species is related to B. longirostris but may be distinguished
from this species by the chaetotaxy of the palpus, having five setae
on the basifemur and tibiotarsus (excluding the end setae) rather
than twelve to fourteen setae on each of these segments. In addi-
tion, hisetosa has two setae on each chelicera rather than one.

Female: Color unknown. Body narrowly ovoid, weakly con-
stricted; length, including gnathosoma, 1100 ijl. Gnathosoma: Length,
277 [a; palpus (fig. 47) short, distal half of the tibiotarsus extending
beyond hypostome; measurements: I, 14 [;.; II, 114 [a; III, 25 \i.; IV,
39 [i.; V, 124 [x; des, 119 [jl; ves, 102 [x. Chelicera normal, nonstriated,
255 [X in length; movable digit with inner surface flattened distally,
fixed digit with preapical tooth and shorter than movable digit;
setae inserted as in Bdella longicornis. Gnathosomal base striated,
buccal cone nonstriated; vh 1-5 equidistant, in longitudinal series,
vh 6 medial between apex and vh 5, vh 1 inserted between palpal
articulations; dorsal hypostomal setae approximately half the length
of vh 1. Dorsal propodosoma (fig. 155): Striae finely broken;
median propodosomals nude, 70 [i in length; eyes separated by
distance equal to three diameters of anterior pair; distance between
anterior sensilla, 60 [x. Dorsal hysterosoma: Setae nude, length of
internal humeral, 51 [x, approximately one half of first interspace;
external humeral, nude, 50 [x in length; sacrals in gently curving
transverse row, clunals in subrectangular arrangement. Anal re-
gion: Border striae parallel; anal setae wanting; one pair of par-
anals; postanals flanking termination of cleft, approximately equal

A Revision of the Mite Family Bdellidae 415

in length to external clunals. Genital region: Each genital plate
with 9 small, equal setae in linear arrangement; 3 pairs of para-
genitals; genital discs small, equidistant; ovipositor with 16 sub-
apical, 4 postmedial setae. Legs (figs. 91, 92): Claws with 5 to 6
lateral rays each, distally alternating with 4 short rays; measure-
ments: tibia I, 70 \l; tarsus I, 170 [k; tibia II, 77 \i.; tarsus II, 170 \i.
Chaetotaxy: coxae I-IV, 6, 3, 6, 2 tactile setae; trochanters I-IV,
1 tactile seta each; basifemora I-IV, 10, 10, 9, 4 tactile setae; telo-
femur I, 7 tactile setae, 1 attenuate sensory seta; telofemora II-IV,
7 tactile setae each; genu I, 5 tactiles, 6 attenuate sensory setae;
genu II, 4 tactiles, 4 attenuate sensory setae; genu III, 5 tactiles,
3 attenuate sensory setae; genu IV, 5 tactiles, 3 attenuate sensory
setae; tibia I, 8 tactiles, 3 attenuate sensory setae, 1 hollow peg,
trichoboth; tibia II, 8 tactiles, 1 attenuate and 1 blunt sensory
seta, trichoboth; tibia III, 10 tactiles, 1 attenuate sensory seta;
tibia IV, 9 tactiles, trichoboth; tarsus I, 13 plumose ventrals arranged
in 2 rows, 5 lateral and 2 dorsal tactile setae, 2 attenuate and 2
blunt sensory setae, 1 peg, dorsoterminals solid, plumose; tarsus II,
14 ventrals as in tarsus I, 5 lateral and 2 dorsal tactile setae, 2 blunt
sensory setae, 1 peg, dorsoterminals solid, plumose; tibia III, ven-
trals, laterals, and dorsoterminals as in tarsus L trichoboth; tarsus
IV, 14 ventrals as in tarsus I, 5 laterals, trichoboth, dt 1 reduced to
1 solid, plumose seta, dt 2, 3 solid, plumose.

Male: Identical to female except in general size and genital
region; length including gnathosoma, 995 [x; genital plates with 8
pairs of setae; amphioid sclerites with 1, 6, 1 setae each.

Holotijpe: Female, 10 miles west of Tuxtla Gutierrez, Chiapas,
Mexico, July 8, 1955, R. E. Beer, under rock.

Allotype: Male, same data as holotype.

Paratypes: Six males and six females, same data as holotype; one
male, one female, 17 miles north of Tehuitzingo, Puebla, Mexico,
July 18, 1955, R. E. Beer, under rocks.

Location of types: The holotype, allotype, and eight paratypes
deposited in the Snow Entomological Museum; two paratypes de-
posited in each of the following: the United States National Mu-
seum, the South Australian Museum, and the British Museum ( Nat-
ural History).

Remarks: The number of pairs of setae on the female genital
plates may be eight, nine, or eleven. When the latter number
occurs, two setae are lateral to the longitudinal series. The am-
phioid sclerites of the male have one or two setae in the anterior

416 The University Science Bulletin

group plus six median setae and one posterior seta. Variation in
general size is slight in the small type series. Because of the two
setae on each chelicera, rather than the usual number of one, this
new species is named hisetosa. Drawings of the holotype.

Cytinae Grandjean, 1938
The members of this subfamily have two pairs of conspicuous
ventral hypostomal setae, maximally three pairs of trichoboths, and
well developed genital tracheae. The palpal tibiotarsus is truncate
and usually slightly longer than the combined length of the genu
and telofemur. The subequal end setae are longer than the palpal
femur. The chelicerae are thickened (Cifta) or normal (Trachy-
molgus) and each bears two setae, one of which is inserted at the
base of the fixed digit. The dorsal propodosoma has lateral pro-
podosomal setae, secondary apodemes well develoed, four eyes
lateral to the unmodified posterior pseudostigmatic organs, and in
Cyta, a fifth, unpaired eye between the anterior sensilla. The
podocephalic canal is an internal tube in Trachymolgus and an
external groove in Cyta (Grandjean, 1938). An unpaired seta
occurs immediately anterior to the genital flaps (between coxae IV
in Cyta spuria, sp. nov).

Genus Cyta von Heyden

Cijta V. Heyden, 1826, Isis von [Lorenz von] Oken, Leipzig (=Encyklo-
padische Zeitsch. vorz. Naturgesch., vergl. Anat. und Phys., vol. 19, no. 6,
p. 608. (Type: Scirus latirostris Hermann,, 1804 [by original designation].)

Amonia Koch, 1836, Deutschlands Crustaceen, Myriapoden und Arachniden.
Ein Beitrag zur Deutschen Fauna von C. L. Koch, Regensburg, fasc. 5,
no. 7. (Type: Amonia cruciata Koch, 1836 [=Cijta latirostris Hermann]
[first included species].)

Ammonia Koch, 1842 {non Bruennich, 1772, Mollusca), Obersicht des Arach-
nidensystems von C. L. Koch, Niirnberg, vol. 3, p. 75. (Type: Amonia
megacephala Koch, 1839 [= Cyta latirostris Hermann] [by original designa-
tion].)

Troglohdella Oudemans, 1937, Kritisch Historisch Overzicht der Acarologie,
Leiden, vol. 3, part C, p. 1228 (new synonym). (Type: Scirus obisium
Gervais, 1841 [^Cyta latirostris Hermann] [by original designation].)

The genus Cyta is related to Trachymolgus, but is distinguished
by thickened chelicerae with massive chelae, and an unpaired me-
dian eye between the anterior sensilla. The distal pair of ventral
hypostomal setae are inserted immediately anterior or posterior to
the suture separating the buccal cone and the strongly developed
lateral lips. The posterior sensilla are widely separated and near
the lateral margins of the dorsal propodosoma. Well developed
lateral apodemes and weakly developed transverse apodemes are
present. There may be three, one, or no pairs of trichoboths. The
podocephalic canal is an external groove (Grandjean, 1938).

A Revision of the Mite Family Bdellidae 417

Troglobdella Oudemans is tentatively placed in synonymy with
Cyta. Oudemans erected this genus for a poorly described and
poorly illustrated species, Scirus ohisium Gervais, 1841. As de-
scribed by Gervais, this species lacks both trichoboths and eyes,
has extremely short palpi and end setae, is light orange to translu-
cent in color, and has the posterior pseudostigmatic organs widely
separated. Except for the apparent lack of eyes, the other distin-
guishing features of Scirus obishim can be found in a proto- or
deutonymph of Cyta latirostris. It is possible to imagine that eyes
could be overlooked because of poor optical equipment or an unsat-
isfactory method of preparation.

Key to Cyta

1. Trichoboths on tibiae I, IV, tarsus III; lateral propodosomals not

approximate to posterior sensilla 2

Trichoboth on tibia IV; lateral propodosomals approximate to
posterior sensilla latirostris p. 417

2. Integument purple; palpal basifemur with more than six setae and

extending well beyond hypostome coerulipes p. 419

Integument not purple; palpal basifemur with less than six setae

and not extending well beyond hypostome spuria p. 421

Cyta latirostris ( Hermann )

(Figs. 30, 60, 164, 172, 178)

Scirus latirostris Hermann, 1804, Mem. Apt., p. 62.

Bdella robustirostris Ewing, 1913, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 112,

plate 7, fig. 3.
Cyta novangliae Jacot, 1939, Occ. Pap. Boston Soc. Nat. Hist., vol. 8, p. 322

( not examined ) ( new synonym ) .

This species is related to Cyta s]mria, sp. nov., but has only one
pair of trichoboths inserted on tibia IV, rather than three pairs
inserted on tibiae I and IV and tarsi III.

Female: Color in life dark red with dark blue-black blotches.
Body short, robust; length, including gnathosoma, 925 |jl (664-
1100 pi). Gnathosoma: Length, 178 [;. (160-221 [x); palpus (fig. 60)
with basifemur not extending beyond hypostome; measurements:
I, 17 ij. (16-20 ij.); II, 124 [X (107-126 ^); III, 29 [;. (24-34 (x); IV,
27 [x (24-34 |x); V, 66 p. (53-68 jx); des, 170 [x (136-196 [x); ves,
131 !x (111-160 [x). Chelicera (fig. 30) inflated, thickened distally,
striated, 173 tx (153-209 jx) in length; inner surface of fixed digit
with 2 distal teeth; setae as figured. Gnathosoma striated; v/i 2
inserted on lateral lips, immediately anterior to articulation of lips;
dorsal hypostomals longer than vh 2. Dorsal propodosoma (fig.
164): Striae finely broken; lateral propodosomals nude, approxi-

14—8920

418 The University Science Bulletin

mately 54 \i (54-80 [x) in length; median propodosomals minutely
pilose, 44 [x (41-71 [i) in length; eyes separated by distance equal
to two and one-half diameters of anterior pair; distance between
anterior sensilla, 110 jji (102-155 [x). Dorsal hysterosoma: Setae
minutely pilose to pilose; length of internal humeral, 44 [i (36-60 [l),
approximately one third to one half of first interspace; external
humeral, 41 [l (37-60 pi) in length; sacrals and clunals in gently
curving transverse rows. Anal region: Border striae parallel; 2
pairs of anal setae; 3 pairs of paranals; postanals wanting. Genital
region: Each genital plate with 8 (7-9) unequal setae in irregular
linear arrangement, anterior 3 setae longer than posterior 6 (5-7)
setae; 6 pairs of paragenital setae, 1 unpaired median seta anterior
to genital plates; genital discs small, equidistant, in central region
of vestibule; ovipositor with 18 subapical, 2 postmedial setae. Legs
(similar to figs. 113, 114) : Claws with one row of minute rays each,
lateral rays wanting; measurements: tibia I, 99 [i. (83-119 \i); tarsus
I, 124 [JL (107-126 [x); tibia II, 61 [;. (51-68 [;.); tarsus II, 85 [l (75-
112 \i). Chaetotaxy: coxae I-IV, 5, 5 (4), 4, 2 tactile setae; trochan-
ters I-IV, 2 (1), 2, 2, 2 (3) tactile setae; basifemora I-IV, 9 (10),
8 (9), 7, 4 tactile setae; telofemora I-IV, 5 (5-7), 6 (7), 4 (4-6),
4 (4-6) tactile setae; genua I-III, 7(6) tactile setae and 1 attenuate
sensory seta each; genu IV, 6 tactile setae; tibia I, 9 (8) tactiles,
1 attenuate and 1 blunt sensory seta, 1 peg; tibia II, 9 (8) tactiles, 1
large and 1 small blunt sensory seta; tibia III, 9(8) tactiles, 1 blunt
sensory seta; tibia IV, 8(7) tactiles, trichoboth; tarsus I, 14 (11-14)
minutely pilose ventral setae arranged in 2 rows, 8 lateral and 2
dorsal tactiles, 2 (1) attenuate and 2 blunt sensory setae, 1 peg,
dt 1, 3 hollow, minutely pilose, dt 2 solid, nude; tarsus II, 11 (11-12)
ventrals, 6 lateral and 2 dorsal tactiles, 1 blunt sensory seta, 1 peg,
dt 1, anterior dt 3 as in tarsus I, dt 2, posterior dt 3 solid, minutely
pilose; tarsus III, 12 (10-13) ventrals, 6 (5) lateral and 1 dorsal tac-
tile seta, dorsoterminals solid, minutely pilose; tarsus IV, 13 (10-13)
ventrals, 1 attenuate sensory seta, dt 1 reduced to one solid, minutely
pilose seta, dt 2, 3 as in tarsus III.

Male: Identical to female; amphioid sclerites as in fig. 172.

Type: Europe.

Location of type: Unknown.

Remarks: Approximately one hundred specimens were examined,
which included material from: Cuba, Jamaica, Mexico (Puebla,
Mexico, San Luis Potosi), the United States (California, Utah,
Idaho, Texas, Arkansas, Kansas, Nebraska, North Dakota, Michigan,

A Revision of the Mite Family Bdellidae 419

Alabama, Tennessee, West Virginia, Maryland, Connecticut),
Alaska (Point Barrow), Iceland, Italy, Germany, and Australia.

Two specimens from Italy have three setae on the palpal genu,
which agrees with Thor's ( 1931 ) description of this species, the
remainder of the specimens have four setae on this segment. The
number of tactile setae on the appendages varies only slightly from
the numbers indicated in the above description, although there
may be four to seven setae on the palpal basifemur. Special sen-
sory setae are constant, except for the occasional absence of one
attenuate sensory seta on tarsus I. Grandjean (1938) reports that
C. latirostris in Europe is actually composed of two species; the
typical species with one pair of trichoboths and another species
lacking trichoboths. Only individuals of the first type were en-
countered in the present study. The description and illustrations
are based on a female from: near Sheep Canyon, Borego State
Park, San Diego Co., California, April 27, 1955, R. O. Schuster,
sycamore and palm litter.

Cijta coerulipes (Duges)
(Figs. 31, 57, 58, 115, 116, 165, 171)

Type: Bdella coerulipes Duges, 1834, Ann. Sci. Nat., ser. 2, vol.
2, p. 45.

This distinctive species is related to C. spuria, sp. no v., but is
easily distinguished by having a purple integument, the palpal
basifemur extending beyond the hypostome, and external sacrals
and clunals longer than the same internal members. In contrast,
C. spuria has a neutral colored integument, the palpal basifemur is
shorter than the hypostome, and the sacrals and clunals are of ap-
proximately equal lengths.

Female: Color in life dark yellow with brown blotches to deep
purple. Body strongly widened at shoulders; length including
gnathosoma, 860 [t. (750-1100 [>.). Cnathosoma: Length, 254 [x (214-
266 |jl); palpus (figs. 57, 58) with elongate basifemur extending
slightly beyond hypostome; measurements: I, 19 [jl (17-20 |x); II,
250 ^ (179-277 ^)- III, 34 ^. (26-36 [j.); IV, 32 ^. (24-34 [x); V, 94 ^ (71-
95 [x); des, 316 [x (292-316 [x); ves, 240 ^. (211-240 ^.). Chelicera (fig.
31) thickened distally, faintly striated, 226 fj. ( 192-238 \).) in length;
fixed digit with broad tooth medially; setae as figured. Gnatho-
somal base and proximal two thirds of buccal cone striated; vh 2
inserted immediately proximal to lateral lips; dorsal hypostomals
approximately equal in length to adornal setae. Dorsal propo-
dosoma (fig. 165): Striae sparsely broken; lateral propodosomals

420 The University Science Bulletin

plumose, 92 [i (66-111 \i) in length; median propodosomals plumose,
129 [X (111-167 [a) in length; eyes separated by distance equal to two
and one half diameters of anterior pair; distance between anterior
sensilla, 112 [x (85-117 n). Dorsal hijsterosoma: Setae plumose;
length of internal humeral, 114 [x (105-145 [x), approximately equal
to first interspace; external humeral, 122 [x (85-170 [x) in length; sa-
crals and clunals in strongly curving transverse rows. Anal region:
Border striae parallel; anal and postanal setae wanting; 3(2) pairs
of paranals. Genital region: Each genital plate with 9 unequal
setae in linear arrangement, anterior 3 longer than posterior 6; 6
(7) pairs of paragenitals; unpaired median seta caudad to coxae
IV; ovipositor with 18 subapical, 2 postmedial setae. Legs (figs.
115, 116): Claws with one row of minute rays each, lateral rays
wanting; measurements: tibia I, 95 [x (78-107 [x); tarsus I, 153 (x
(128-160 !x); tibia II, 83 (x (60-95 [x); tarsus II, 136 [x (119-146 (x).
Chaetotaxy: coxae I-IV, 5 (4), 4 (5), 5 (6), 4 (3-5) tactile setae;
trochanters I-IV, 2 tactile setae each; basifemora I-IV, 7 (8), 7
(5-8), 7 (6-8), 5 (4) tactile setae; telofemora I-IV, 5 (5-7), 5
(4-6), 4 (5), 4 (3) tactile setae; genu I, 4 (3) tactiles, 2 attenuate
sensory setae; genu II, 4 (5) tactiles, 1 attenuate sensory seta; genu
III, 5 (4-6) tactiles, 1 attenuate sensory seta; genu IV, 6 (5) tac-
tiles, 1 attenuate sensory seta; tibia I, 8 (8-10) tactiles, 2 attenuate
sensory setae, 1 peg, trichoboth; tibia II, 9 (8) tactiles, 1 attenuate
and 1 blunt sensory seta; tibia III, 9 (8-10) tactiles, 1 attenuate
sensory seta; tibia IV, 10 (9) tactiles, trichoboth; tarsus I, 12 (11-
13) pilose ventrals arranged in two rows, 8 (7) lateral and 2 dor-
sal tactiles, 2(1) attenuate and 2 blunt sensory setae, dorsotermi-
nals solid, pilose; tarsus II, 10 (11) ventrals, 7 (8) lateral and 2
dorsal tactiles, 1 attenuate and 1 blunt sensory seta, dorsoterminals
as in tarsus I; tarsus III, 12 ventrals, 8 laterals, trichoboth, dorso-
terminals as in tarsus I; tarsus IV, 12 (11) ventrals, 6 laterals, 1
attenuate sensory seta, tricohboth wanting, dt 1 reduced to one
solid, pilose seta, dt 2, 3 solid, pilose.

Male: Identical to female; amphioid sclerites each with 9 (8)
setae (fig. 171).

Type: Europe.

Location of type: Unknown.

Remarks: Approximately 200 slides were examined, which in-
cluded specimens from: Panama, Haiti, Cuba, Mexico (San Luis
Potosi, Guerrero, Tamaulipas, Hidalgo), the United States (Colo-
rado, California, Texas), Alaska (Umiat) and Sudan Bor.

A Revision of the Mite Family Bdellidae 421

Slight, but constant variation in the widely separated localities
indicates that C. coenilipes may be a group of closely related spe-
cies or subspecies. Extreme variation is found in the forms of the
palpal setae. In southeastern Texas and northeastern Mexico, the
small tactile setae are strongly plumed (fig. 58), a condition not
found in other geographical areas. The attenuate sensory setae on
tibia I undergo considerable migration and reduction in numbers,
for example, the Cuban specimens have the two approximate at-
tenuate sensory setae near the distal margin of the segment rather
than the proximal margin (fig. 115). Specimens from Panama
have three sensory setae in a linear arrangement, while those from
Colorado and Sudan Bor have only two attenuate sensory setae.

The striae on the dorsal propodosoma are usually almost con-
tinuous, but in the insular material, the striae are finely broken, re-
sembling those found in Cyta latirostris (fig. 164).

The redescriptions and illustrations are based on a male and a
female from: five miles east of Ciudad del Maiz, San Luis Potosi,
Mexico, August 23, 1954, W. T. Atyeo, ground litter, altitude 4,700
feet. An illustration of plumose setae on the palpus was prepared
from a male from: ten miles north of Edinburg, Texas, April 9,
1954, R. E. Beer, under rock.

Ctfta spuria, sp. nov.
(Figs. 11, 59, 113, 114, 167, 173)

In size and shape, this species closely resembles Cyta latirostris,
but may be distinguished by the presence of trichoboths on tibiae
I, IV, and tarsus III rather than trichoboths only on tibia IV.

Female: Color in life unknown. Body short, robust, weakly
constricted; length, including gnathosoma, 810 a. Gnathosoma:
Length, 194 [jl; palpus (fig. 59) with basifemur-lelofemur articula-
tion indistinct; measurements: I, 14 pi.; II plus III, 138 \i; IV, 22 [/.;
V, 61 [x; des, 179 [i: ves, 136 [jl. Chelicera inflated, striated, 158 \i.
in length; fixed digit smooth, longer than movable digit (fig. 11);
distal seta inserted above proximal limit of movable digit, proximal
seta as in C. latirostris, except extending beyond articulation of
chela. Gnathosomal base and proximal two thirds of buccal cone
striated; vh 2 immediately proximal to lateral lips; dorsal hyposto-
mal setae equal in length to vh 2. Dorsal propodosoma (fig. 167):
Striae very finely broken; lateral propodosomals plumose, 77 \l in
length; median propodosomals plumose, 78 [i in length; eyes sep-
arated by distance equal to two and one half diameters of anterior
pair; distance between anterior sensilla, 97 [x. Dorsal hysterosoma:

422 The University Science Bulletin

Setae plumose; length of internal humeral, 73 [x, approximately three
fourths of first interspace; external humeral, 77 \j. in length; sacrals
and clunals in strongly curving transverse rows. Anal region: Bor-
der striae parallel; anal and postanal setae wanting; 3 pairs of
paranals. Genital region: Each genital plate with 9 unequal setae
in linear arrangement, anterior 3 longer than posterior 6; 9 pairs
of paragenitals; 1 median unpaired seta between coxae IV; oviposi-
tor with 20 setae of undeterminable position. Legs (figs. 113, 114) :
Claws with row of minute rays each, lateral rays wanting; measure-
ments: tibia I, 73 [a; tarsus I, 128 [a; tibia II, 68 pi; tarsus II, 133 ^.
Chaetotaxy: coxae I-IV, 5 (of which 1 longer than femur II), 1,
5, 2 tactile setae; trochanters I-IV, 2, 2, 2, 1 tactile setae; basifemora
I-IV, 8, 8, 7, 5 tactile setae; telofemora I-IV, 5, 5, 4, 4 tactile setae;
genua I-IV, each with 4 tactile setae and duplex seta with microseta
in form of short, hollow peg; tibia I, 8 tactiles, 2 attenuate sensory
setae, 1 peg, trichoboth; tibia II, 8 tactiles, 1 attenuate and 1 blunt
sensory seta; tibia III, 8 tactiles, 1 attenuate sensory seta; tibia IV,
9 tactiles, trichoboth; tarsus I, 12 minutely pilose ventrals in 2 rows,
8 lateral and 2 dorsal tactile setae, 1 attenuate and 2 blunt sensory
setae, dorsoterminals solid, minutely pilose; tarsus II, similar to tar-
sus I, except lacking 1 attenuate and 1 blunt sensory seta; tarsus III,
ventrals, laterals and dorsoterminals as in tarsus I, trichoboth; tarsus
IV, ventrals and dorsoterminals as in tarsus I, 6 lateral tactile setae,
1 attenuate sensory seta.

Male: Identical to female, except slightly smaller in general
size; total length, 710 [j.; amphioid sclerites as in fig. 173.

Holotype: Male, 10 miles east of Xihtla, San Luis Potosi, Mexico,
July 25, 1954, Warren T. Atyeo, beating bamboo.

Allotype: Female, same data as holotype.

Paratypes: Six males, nine females, same data as holotype; one
male, one female, 12 miles east of Xilitla, San Luis Potosi, Mexico,
June 20, 1955, R. E. Beer, on bamboo; one male, 10 miles east of
Tuxtla Gutierrez, Chiapas, Mexico, July 8, 1955, R. E. Beer, under
rock; one female, Huatusco, Veracruz, Mexico, at Brownsville,
Texas, August 22, 1949, with orchid plants.

Location of type: The holotype, allotype, and thirteen paratypes
deposited at the Snow Entomological Museum; two paratypes de-
posited at each of the following: the United States National
Museum, the British Museum (Natural History), and the South
Australian Museum.

A Revision of the Mite Family Bdellidae 423

Remarks: The proximal setae on the appendages tend to be mi-
nutely pilose while the distal setae tend to be pilose. Except for
the slight variation in the number of tactile setae on the leg seg-
ments, the other features of this species are very stable. Total
length, including gnathosoma, approximately 699 \). to 800 [x. Draw-
ings of the holotype.

Genus Trachymolgus Berlese

Trachymolgus Berlese, 1923, Redia, vol. 15, p. 242. (Type: Bdella niger-
rima Canestrini and Fanzago, 1876 [by original designation].)

Although related to Cyta, this European genus has two unique
features. The integument of the idiosoma is heavily sclerotized
and reticulated, and instead of round genital tracheae, this group is
characterized by platytracheae which expand near their termina-
tions over coxae II (Grandjean, 1938). The chelicerae are normal
and the small chelae are dentate. The cheliceral setae are arranged
as in Cyta, that is, the distal seta is inserted at the base of the fixed
digit. The palpal tibiotarsus is not strongly shortened nor widened
distally, but elongated as in Biscirus and Monotrichobdella. Two
pairs of widely separated eyes and four pairs of setae occur on the
dorsal propodosoma; the podocephalic canal is an internal tube
(Grandjean, 1938). Three pairs of trichoboths are inserted on the
legs. Specimens of this genus were not available for examination.

Spinibdellinae Grandjean, 1938

The Spinibdellinae is distinguished by two pairs of conspicuous
ventral hypostomal setae, four pairs of trichoboths, and well
developed genital tracheae. The palpal tibiotarsus is truncate and
shorter than the combined length of the genu and telofemur in
Spinibdella or cylindrical and longer than the combined segments
in Bi&cirus and Monotrichobdella. The subequal end setae are as
long, or longer than the palpal femur. Elongated chelicerae bear
two setae inserted on the distal two thirds and small, needlelike
chelae. The dorsal propodosoma with or without lateral propo-
dosomal setae; secondary apodemes absent or poorly developed;
two or four eyes lateral to the unmodified posterior pseudostigmatic
organs; and the propocephalic canal is an internal tube (Grandjean,
1938 ) . If present, unpaired setae on the venter of the hysterosoma
are between the coxae, not immediately anterior to the genital flaps.

The three genera comprising this subfamily are similar. They
are separated primarily by the shape of the palpal tibiotarsus, the
number of end setae, and the presence or absence of the lateral
propodosomal setae.

424 The University Science Bulletin

Genus Spinibdella Sig Thor

Spinibdella Thor, 1930, Zool. Anz., vol. 92, no. 1, p. 22. (Type: Spinibdella
reducta Thor, 1930 [by original designation].)

Spinibdella is closely related to Biscirus, but has a short and
truncated palpal tibiotarsus, and the lateral propodosomal setae
are present. The palpal genu has three or four setae, and the basi-
femur has more than two setae. The cheliceral setae may be minute
or of medium length, with the distal seta not extending beyond the
tips of the chelae. Striation patterns on the dorsum of the propo-
dosoma appear to be specific as do the modified setae on the am-
phioid sclerites of the males.

Key to the Species of Spinibdella

1. Palpus much shorter than the hypostome; cheliceral setae mi-
nute tenuirostris p. 424

Palpus longer than hypostome; cheliceral setae conspicuous 2

2. Palpal tibiotarsus approximately tlie same length and diameter of

palpal genu corticis p. 426

Palpal tibiotarsus longer and thicker than the genu 3

3. Blunt sensory seta on tibia II deeply recessed 4

Blunt sensory seta on tibia II not recessed 5

4. One pair of eyes; internal humerals half of first interspaces

depressa p. 428
Two pairs of eyes; internal humerals equal to first interspaces

hifurcata p. 430

5. Internal humerals equal to first interspaces; tibia II with one blunt

sensory seta cronini p. 432

Internal humerals half of first interspaces; tibia II with one blunt

and one attenuate sensory seta ornata p. 434

Spinibdella tenuirostris (Ewing), new combination

(Figs. 32, 61, 123, 124, 157)

Bdella tenuirostris Ewing, 1914, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 149.
Spinibdella wilsoni Jacot, 1938, Psyche, vol. 45, no. 2-3, pp. 129-130 (new
synonym ) .

This species is closely related to, or conspecific with Spinibdella
reducta Thor, 1930, and possibly Spinibdella lignicola (Berlese),
1892 {=z Bdella lignicola Berlese), but the inadequate descriptions
make it impossible to determine the true synonymies of this species.
Both tenuirostris and reducta have the gnathosoma considerably
longer than the palpus, and both have minute cheliceral setae.
Berlese's species with "setis mandibularwn deficientibus vel minimis,
palpisque cortioribus" can not be evaluated with certainty.

Female: Color unknown. Body pear-shaped, weakly constricted;
length, including gnathosoma, 1100 [i. Gnathosoma: Length, 330 [jl;

A Revision of the Mite Family Bdellidae 425

palpus (fig. 61) considerably shorter than hypostome; measure-
ments: I, 15 [x; II plus III, 153 [x; IV, 20 [>.; V, 54 [x; des, 185 [X;
ves, 160 [X. Chehcera nonstriated, 306 [>. in length; setae small, each
9 [x in length, inserted on distal half of chelicera (fig. 32). Gnatho-
soma striated proximal to ventral setae; dorsal hypostomal setae
wanting. Dorsal propodosoma ( fig. 157 ) : Striae coarsely broken;
lateral propodosomals nude, 46 [x in length; median propodosomals
nude, 60 [x in length; eyes subequal, separated by distance equal to
2 diameters of smaller, posterior pair, interval between eyes with
transverse striae; distance between anterior sensilla, 90 [x. Dorsal
hijsterosoma: Setae nude; length of internal humeral, 83 [x, approxi-
mately three fourths of first interspace; external humeral, 83 |x in
length; sacrals and clunals in gently curving transverse rows. Anal
region: Border striae parallel; 3 pairs of anal setae; 3 pairs of par-
anals; postanals distant from termination of cleft, shorter than
clunals. Genital region: Each genital plate with 8 equal setae in
linear arrangement; 10 pairs of paragenitals, anterior pair between
coxae IV; ovipositor with 18 setae of interminable position. Legs
(figs. 123, 124): Claws small, shorter than pretarsus, with 4-5
lateral rays; measurements: tibia I, 97 [x; tarsus I, 128 [x; tibia II,
78 [x; tarsus II, 119 [x. Chaetotaxy: coxae I-IV, 5, 5, 5, 3 tactile setae;,
trochanters I-IV, 1, 1, 2, 1 tactile setae; basifemora I-IV, 11, 10, 9,
4 tactile setae; telofemora I-IV, 11, 10, 8, 6 tactile setae; genu I, 7
tactiles, 5 attenuate sensory setae, 1 small, solid seta; genu II, 7
tactiles and 1 attenuate sensory seta; genua III-IV, 7 tactiles, 1
attenuate sensory seta each; tibia I, 15 tactiles, 8 attenuate sen-
sory setae, 1 attenuate peg, trichoboth; tibiae II-III, 13 tactiles, 1
attenuate and 1 blunt sensory seta; tibia IV, 12 tactiles, trichoboth;
tarsus I, 14 plumose ventrals arranged in 2 rows, 9 lateral and 2
dorsal tactiles, 3 attenuate and 2 blunt sensory setae, 1 capitate
peg, dt 1 hollow, nude, dt 2, S hollow; plumose; tarsus II, 12 plumose
ventrals, 8 lateral and 2 dorsal tactile setae, 2 blunt sensory setae,
1 peg, dt 1 solid, nude, dt 2, 3 solid, plumose; tarsus III, 12 plumose
ventrals, 9 laterals, trichoboth, dorsoterminals as in tarsus II; tarsus
IV, 12 plumose ventrals, 7 laterals, trichoboth, attenuate sensory seta
posterolateral of trichoboth, dorsoterminals as in tarsus II.

Male: Unknown.

Type: Female, Bdella tenuirostris, Xenia, Ohio, September 14,
1910, H. E. Ewing, under stones.

Location of tifjje: The United States National Museum.

426 The University Science Bulletin

Remarks: Three female cotypes of Spinibdella tvilsoni Jacot were
examined; these were collected 7 miles from south point of North
Beach, St. Augustine, Florida, March 7, 1928, E. F, Grossman, leaf
litter of Tamola littoralis on shore bay. These cotypes are further
identified by E. F. Grossman's code numbers: two females, G34Bdl
and one female, G34Bd3. In addition to the types, specimens were
examined from Florida, Arkansas, Kansas, North Garolina, Vermont,
Michigan, and Galifornia,

This is a highly variable species, and although subspeciation is
probable, no correlations of characters can be made with the speci-
mens available for study. Extensive variation occurs in the special
sensory setae on tibia I, the chaetotaxy of the palpus, the number
of setae in the genital region, and the general size. The California
and Arkansas specimens have only one long attenuate sensory seta
distal to the trichoboth on tibia I, whereas the specimens from other
locales have two setae in this position. The proximal sensory setae
of tibia I vary from five to six in number, and in the specimens from
Florida, North Carolina, Arkansas and California, they are approxi-
mately one fourth the length of the tibia. In the Kansas specimens
and the tvilsoni cotypes, the same sensory setae are almost one half
the tibial length.

Variation in the number of setae on the palpus is confined to the
basifemur and genu. The Kansas specimens have five setae on the
basifemur, while the remaining specimens have seven setae; the
genua of individuals from Kansas, Michigan, and Florida (Jacot's
cotypes ) have four setae, the other specimens have only three. The
genital setae vary from five to eight pairs, and there may be as
many as two unpaired paragenital setae between the posterior
coxae. Total length varies from 880 \j. to 1420 \j.. Drawings of
Ewing's type.

Spinibdella corticis (Ewing), new combination
(Figs. 33, 66, 127, 128, 156)
Bdella corticis Ewing, 1909, Can. Ent., vol. 41, no. 4, p. 122.

Although related to Spinibdella biftircata, sp. nov., this species
is distinctive in having the palpal tibiotarsus approximately equal
in length and diameter to the palpal genu. S. biftircata, as well as
the other known species of this genus, has the palpal tibiotarsus
considerably larger than the genu.

Female: Color unknown. Body pear-shaped; length, including
gnathosoma, 1243 p.. Gnathosoma: Length, 391 iji; palpus (fig. 66)
with genu longer than tibiotarsus; measurements: I, 25 [x; II, 291 [x;

A Revision of the Mite Family Bdellidae 427

III, 33 [l; IV, 58 ii; V, 50 i).; des, 411 ^■, ves, 349 [j.. Chelicera (fig. 33)
striated, 347 ^ in length; setae as figured. Gnathosoma striated;
dorsal hypostomal setae conspicuous, 48 [j. in length. Dorsal propo-
dosoma (fig. 156): Striae sparsely broken; lateral propodosomals
with minute branchings, 84 \). in length; median propodosomals
branched, 126 [j. in length; eyes separated by distance equal to
diameter of anterior pair, interval between eyes with longitudinal
striae; distance between anterior sensilla, 68 [x. Dorsal hystero-
soma: Setae branched; length of internal humeral, 118 pi., approxi-
mately the length of the first interspace; external humeral, 149 \t. in
length; sacrals in strongly curving transverse row, clunals in sub-
trapazoidal arrangement. Anal region: Border striae parallel, 2
pairs of anal setae; 1 pair of paranals; postanals flanking anal cleft,
shorter than clunals. Genital region: Each genital plate with 10
long setae in hnear arrangement; 28 pairs of paragenitals, anterior
4 pairs between coxae IV; median, unpaired setae wanting; genital
discs small, distant; ovipositor short, with 14 pairs of subapical
setae. Legs (figs. 127, 128): Claws small, shorter than pretarsus,
each with one row of minute lateral rays; measurements: tibia I,
99 [x; tarsus I, 126 i;,; tibia II, 99 \i; tarsus II, 137 (x. Chaetotaxy:
coxae I-IV, 10, 8, 8, 7 tactile setae, many equal in length to basife-
mur I; trochanters I-IV, 1, 1, 2, 1 tactile setae; basifemora I-IV, 13,
12, 7, 4 tactile setae; telofemora I-IV, 8, 9, 6, 6 tactile setae; genua
I-III, 6 tactiles and 1 duplex seta each; genu IV, 6 tactiles; tibia I,
18 tactiles, 2 attenuate sensory setae, 1 peg, trichoboth; tibia II,
18 tactiles, 1 attenuate and 1 blunt sensory seta; tibia III, 15 tactiles,
1 attenuate sensory seta; tibia IV, 14 tactiles, trichoboth; tarsus I,
16 ventrals, 9 lateral and 2 dorsal tactiles, 1 attenuate and 2 blunt
sensory setae, dt 1, 3 hollow, nude, dt 2 solid, nude; tarsus II, 17
ventral, 8 lateral and 2 dorsal tactile setae, 1 attenuate and 1 blunt
sensory seta, dt 2, posterior dt 1 solid, nude, dt 3, anterior dt 1 hol-
low, nude; tarsus III, 20 ventrals, 8 laterals, trichoboth, dt 1, 2
solid, nude, dt 3 hollow, nude; tarsus IV, 19 ventrals, 6 laterals,
trichoboth, setae of dt 1 obliquely arranged, dt 1, 2 solid, nude, dt
3 hollow, nude.

Male: Unknown.

Type: Female, Urbana, iHinois, July 2, 1908, H. E. Ewing, imder
bark of cottonwood tree.

Location of type: The United States National Museum.

Remarks: The type specimen is incomplete and is 966 [i in length,
slightly smaller than the specimen used for the redescription. Only

428 The University Science Bulletin

the type and five additional specimens were available for study;
these latter specimens are identified by the following data: One
female, six miles north of Chila, Puebla, Mexico, July 17, 1955, R. E.
Beer, under stone; one female, Guatemala, at Brownsville, Texas,
February 27, 1946, on Odontoglossinn sp.; one female, south end
of Cedar Mountains, Tooele Co., Utah, July 13, 1953, D. Porter;
one female, Halsey, Thomas Co., Nebraska, 1954, W. F. Rapp, Jr.,
ex: duff; one female, Monte Alban, Oaxaca, Mexico, July 5, 1955,
R. E. Beer, under rock.

In all the specimens, except the Utah female, the special sen-
sory setae of tibia I are grouped in a small area distal to the tricho-
both; in the Utah specimen, one of the attenuate sensory setae has
migrated laterad of the trichoboth. The tactile setae of the three
females from the more southern localities tend to have a greater
diameter and to be more plumose than the same setae of the
United States specimens.

The redescription and illustrations are based primarily on the
female from Monte Alban, Oaxaca, Mexico.

Spinibdella depressa (Ewing), new combination

(Figs. 62, 119, 120, 158, 175)

Bdella depressa Ewing, 1909, Can. Ent., vol. 41, no. 4, p. 125.

Bdella virgata Ewing, 1910, Univ. Stud., Univ. Illinois, vol. 3, no. 6, p. 70

(new synonym).
Bdella chapultepecensis Baker and Balock, 1944, Proc. Ent. Soc. Washington,

vol. 46, no. 7, p. 177 ( new synonym ) .
Bdella rio-lermensis Baker and Balock, 1944, Proc. Ent. Soc. Washington, vol.

46, no. 7, p. 178 ( new synonym ) .

Spinibdella depressa is closely related to S. hifiircata, sp. nov., but
can be distinguished by the positions of the lateral propodosomal
setae and the number of eyes. S. depressa has the lateral propo-
dosomals approximate to the anterior sensilla rather than midway
between the anterior and posterior sensilla, and has one pair of eyes
rather than two pairs.

Female: Color in life light red with large black spots at postero-
lateral margins of the dorsal propodosoma. Body narrow, strongly
constricted; length, including gnathosoma, 753 t;.. Gnafhosoma:
Length, 195 [x; palpus (fig. 62) short, one half of tibiotarsus ex-
tending beyond hypostome; measurements: I, 14 [a; II plus III,
136 tx; IV, 17 [i; V, 40 \).; des, 168 [i; ves, 119 [x. Chehcera finely
striated, 182 [x in length; setae inserted as in fig. 33. Gnathosoma
striated; dorsal hypostomal setae small, 17 [x in length. Dorsal
propodosoma (fig. 158): Striae sparsely broken; lateral propodo-
somals nude, 34 ix in length; median propodosomals plumose, 39 [x

A Revision of the Mite Family Bdellidae 429

in length; posterior pair of eyes wanting, former position indicated
by striae forming teardrop pattern; distance between anterior sen-
silla, 45 [x. Dorsal hijsterosomo: Setae plumose; length of internal
humeral, 34 \i, less than one half of first interspace; external humeral,
34 [i in length; sacrals and clunals in subtrapazoidal arrangement,
external clunals lateral to termination of cleft. Anal region: Border
striae parallel; 1 pair of anal setae; paranals wanting; postanals at
posterior two thirds of cleft, anterior to external clunals. Genital
region: Each genital plate with 9 equal genital setae in linear ar-
rangement; 11 pairs of paragenitals; 4 median unpaired para-
genitals, anterior unpaired seta between coxae III; genital discs
small, 2 pairs in anterior half of vestibule, 1 pair in caudal fourth;
ovipositor with 12 subapical and 6 postmedial setae. Legs (figs.
119, 120): Claws small, unadorned, shorter than pretarsus; meas-
urements: tibia I, 52 [7.; tarsus I, 65 [x; tibia II, 49 [jl; tarsus II,
65 [JL. Chaetotaxy: coxae I-IV, 9, 8, 7, 5 tactile setae; trochanters
I-IV, 1, 1, 2, 1 tactile setae; basifemora I-IV, 7, 8, 7, 3 tactile setae;
telofemora I-IV, 5, 5, 4, 5 tactiles; genua I-III, 6 tactiles and 1 duplex
seta each; genu IV, 7 tactiles; tibia I, 11 tactiles, 1 blunt and 1
attenuate sensory seta, 1 attenuate peg, trichoboth; tibia II, 7
tactiles, 1 recessed blunt sensory seta; tibia III, 12 tactiles, 1 at-
tenuate sensory seta; tibia IV, 11 tactiles, trichoboth; tarsus I, 13
ventrals, 4 lateral and 2 dorsal tactiles, 1 attenuate and 2 blunt
sensory setae, dt 1, 3 hollow, nude, dt 2 solid, nude; tarsus II, as
in tarsus I except lacking 1 attenuate sensory seta and posterior
dt 1 sohd, nude; tarsi III-IV, 13 ventrals, 4 laterals, trichoboth,
dt 1,2 solid, nude, dt 3 hollow, nude.

Male: Identical to female except in genital region: 12-13 pairs
of genital setae; amphioid sclerites as in fig. 175.

Type: Female, Areola, Illinois, July 4, 1908, H. E. Ewing, under
bark.

Location of type: The United States National Museum.

Remarks: The following types were also available for study:
Bdella virgafa Ewing, female, Mahomet, Illinois, April 17, 1908,
H. E. Ewing, under bark; Bdella rio-lermensis Baker and Balock,
female, Rio Lerma, Mexico-Toluca Highway, Mexico, January 24,
1943, E. W. Baker, lichens; Bdella chapuJtepecensis Baker and
Balock, male, Chapultepec Park, Mexico, Distrito Federal, Mexico,
March 16, 1943, E. W. Baker, lichens. Additional males and females,
totaling thirty-six specimens, were examined from Mexico (San

430 The University Science Bulletin

Luis Potosi, Mexico, Morelos ) and the United States ( Texas, Mary-
land, Arkansas, Kansas, Illinois, New Jersey, Connecticut).

Of the widely distributed species, this group exhibits the least
amount of variation. Total length varies from 590 [i to 780 [l, and
only occasionally do the tactile setae of the legs vary from one
more, to one less, than the numbers indicated in the description.
Drawings of a male from: Palmetto State Park, Texas, April 4,
1954, R. E. Beer, Spanish moss.

Spinibdella hifurcata, sp. nov.
(Figs. 1, 2, 63, 117, 118, 162, 174)

Tliis species is closely related to Spinibdella ornata, sp. nov., but
is distinguished by having one blunt sensory seta deeply recessed
on tibia II rather than one blunt and one attenuate sensory seta.
The amphioid sclerites of the males are unique in both species
(figs. 174, 176).

Female: Color unknown. Body (figs. 1, 2) narrow, strongly
constricted; length, including gnathosoma, 702 \}.. Gnathosoma:
Length, 178 [).; palpus (fig. 63) short, tibiotarsus extending beyond
hypostome; measurements: I, 9 [i; II plus III, 125 [i; IV, 18 [a; V,
31 [x; des, 202 [x; ves, 135 [x. Chelicera striated, 167 [i in length;
setae as in fig. 33. Gnathosoma striated; dorsal hypostomal setae
wanting. Dorsal propodosoma (figs. 1, 162) : Striae sparsely broken;
lateral propodosomals nude, 48 [i in length; median propodosomals
finely plumose, 51 [i in length; eyes separated by distance equal to
diameter of anterior pair, interval between eyes with longitudinal
striae; distance between anterior sensilla, 44 [x. Dorsal hijsterosoma:
Setae finely plumose; length of internal humeral, 53 pi, approxi-
mately equal to first interspace; external humeral, 55 \i in length;
sacrals in subtrapazoidal arrangement, external clunals wanting.
Anal region (fig. 2): Border striae parallel; 2 pairs of anal setae;
1 pair of paranals; postanals flanking termination of cleft, shorter
than clunals. Genital region (fig. 2): Each genital plate with 9
equal setae arranged in two rows; 15 pairs of paragenitals, anterior
3 pairs between coxae IV; median unpaired seta between coxae III
wanting; genital discs small, distant; ovipositor with 12 subapical,
6 postmedial setae. Legs (figs. 117, 118): Claws unadorned, small,
shorter than pretarsus; measurements: tibia I, 55 [jl; tarsus I, 71 pi;
tibia II, 48 ix; tarsus II, 70 [j.. Chaetotaxy: coxae I-IV, 9, 8, 7, 6
tactile setae; trochanters I-IV, 1, 1, 2, 1 tactile setae; basifemora
I-IV, 8, 8, 7, 3 tactile setae; telofemora I-IV, 6, 5, 4, 5 tactile setae;
genu I, 5 tactiles, 1 duplex seta; genua II-III, 6 tactiles and 1 duplex

A Revision of the Mite Family Bdellidae 431

seta each; genu IV, 7 tactiles; tibia I, 14 tactiles, 1 attenuate and

1 blunt sensory seta, trichoboth; tibia II, 13 tactiles, 1 recessed,
blunt sensory seta; tibia III, 15 tactiles, 1 attenuate sensory seta;
tibia IV, 15 tactiles, trichoboth; tarsus I, 17 tactiles, 5 lateral and

2 dorsal tactiles, 1 attenuate and 2 blunt sensory setae, dt 1, 3 hol-
low, nude, dt 2 solid, nude; tarsus II, 15 ventrals, 5 lateral and 2
dorsal tactiles, 2 blunt sensory setae, dt 2, anterior dt 1 solid, nude,
dt 3, posterior dt 1 hollow, nude; tarsus III, 18 ventrals, 8 laterals,
trichoboth, dt 1, 2 solid, nude, dt 3 hollow, nude; tarsus IV, 17
ventrals, 6 laterals, trichoboth, dorsoterminals as in tarsus III.

Male: Identical to female except in genital region; each genital
plate with 14 setae; amphioid sclerites as in fig. 174.

Holotijpe: Male, 10 miles west of Tuxtla Gutierrez, Chiapas,
Mexico, July 8, 1955, R. E. Beer, under rock.

Allotype: Female, 16 miles north of Juchitan, Oaxaca, Mexico,
July 2, 1955, R. E. Beer, free hving on various plants.

Paratypes: Six males, five females, same data as the holotype;
one female, same data as allotype; two males, 8 miles south of
Nochistlan, Oaxaca, Mexico, June 30, 1955, R. E. Beer, under rock;
two males, Monte Alban, Oaxaca, Mexico, July 17, 1955, R. E. Beer,
under rock; one male, same data as the preceding, except collected
July 15, 1955; one female, Huajuapam de Leon, Oaxaca, Mexico,
July 17, 1955, R. E. Beer, under rock; one male, 17 miles north of
Tehuitzingo, Puebla, Mexico, July 17, 1955, R. E. Beer, under rock;
one male, one female, 3 miles north of Manzanillo, Michoacan,
Mexico, July 26, 1955, R. E. Beer, on beach under coconut hull;
one female, Mexico, at Brownsville, Texas, July 27, 1952, Smith,
with pineapple fruit; one male, one female, 10 miles north of
Edinburg, Texas, April 4, 1954, R. E. Beer, under log; two females,
Corpus Christi State Park, Texas, August 6, 1955, R. E. Beer, under
rock.

Location of types: The holotype, allotype, and twenty paratypes
are deposited in the Snow Entomological Museum; two paratypes
are deposited in each of the following: the United States National
Museum, the South Australian Museum, and the British Museum
(Natural History).

Remarks: The normally bifurcate seta on each of the amphioid
sclerites (fig. 174) is undivided in one specimen, the resultant seta
is unilaterally plumose. The genital plates of the female bear eight
to ten pairs of setae, those of the males, thirteen to fourteen pairs.
Other than these variations in the genital region and in total length.

432 The University Science Bulletin

which varies from 617 [l to 930 [jl, the characteristic structures of
this species are constant. This species is named bifurcata because
of the unique structure of a median seta on each of the male
amphioid sclerites. Drawings of the holotype.

Spinibdella cronini (Baker and Balock), new combination

(Figs. 65, 125, 126, 159, 160)

Bdella cronini Baker and Balock, 1944, Proc. Ent. Soc. Wasliington, vol. 46,
no. 7 p. 178.

The longitudinally or obliquely directed striae in the center of
the propodosomal shield distinguishes this species from the closely
related Spiiiibdella ornata, sp. nov., in which the striae are horizon-
tally directed.

Female: Color in life pale red with darker markings on dorso-
lateral propodosoma. Body elongate, strongly constricted; length,
including gnathosoma, 753[x (717-1257 \i.). Gnathosoma: Length,
211 [X (194-247 [jl); palpus (fig. 65) short, tibiotarsus extending
beyond hypostome; measurements: I, 10 [a (9-15 [j.); II plus III,
152 [i (126-175 [x); IV, 22 jx (21-27 jx); V, 36 [x (33-43 jx); des,
201 I). (170-254 ix); ves, 131 |x (122-167 jx). Chehcera striated,
196 [X (177-226 [x) in length; setae inserted as in fig. 33. Gnatho-
soma striated; dorsal hypostomal setae small, approximately 16 [x
in length. Dorsal propodosoma (figs. 159, 160): Striae sparsely
broken; lateral propodosomals thickened, nude (to slightly plu-
mose), 49 [X (47-98 [x) in length; median propodosomals thickened,
nude to plumose, 74 [x (53-82 [x) in length; eyes separated by dis-
tance equal to 2 diameters of anterior pair, interval between eyes
with longitudinal striae; distance betwen anterior sensilla, 41 [x
(41-76 [x). Dorsal hysterosoma: Setae thickened, nude (to plu-
mose); length of internal humeral, 76 [x (54-118 [x), approximately
equal to first interspace; external humeral, 69 [x (49-77 [x) in length;
sacrals and clunals in subtrapazoidal arrangement. Anal region:
Border striae parallel; 2 pairs of anal setae; paranals wanting; post-
anals flanking termination of cleft, shorter than clunals. Genital
region: Each genital plate with 13 (13-16) equal, basally con-
stricted setae in irregular linear arrangement; 21 (21-23) pairs of
paragenital setae, anterior pairs between coxae III and IV; 2 (2-3)
median, unpaired paragenital setae, anterior unpaired seta between
coxae III; genital discs small, 2 pairs proximate in anterior half
of vestibule, 1 pair in caudal one fourth; ovipositor short, seen
only in axial view, 6 (5) pairs of setae anterior, 3 pairs posterior
of center. Legs (figs. 125, 126): Claws nude, small, shorter than

A Revision of the Mite Family Bdelledae 433

pretarsus; measurements: tibia I, 58 \i (49-72 [jl); tarsus I, 77 [i
(65-97 ijl); tibia II, 50 [i. (45-65 [l); tarsus II, 76 [i (63-93 [x).
Chaetotaxy: coxae I-IV, 7 (6), 7 (6-8), 7 (8), 6 (5) tactile setae;
trochanters I-IV, 1, 1, 2, 1 tactile setae; basifemora I-IV, 7, 7 (8),
5, 3 tactile setae; telofemora I-IV, 5, 5, 4, 4 (3) tactiles; genua
I-IV, 5, 5, 5, 6 tactiles; tibia I, 13 (12) tactiles, 1 attenuate sensory
seta, 1 peg, trichobotli; tibia II, 12 (13) tactiles, 1 blunt sensory
seta; tibia III, 11 (12) tactiles, 1 attenuate sensory seta; tibia IV, 14
tactiles, trichoboth; tarsus I, 12 (11) ventrals, 6 (7) lateral and 2
dorsal tactile setae, 2 blunt sensory setae, dt 1, 2 solid, nude, dt 3
solid, nude (one or both may be hollow, nude); tarsus II, 11 (10)
ventrals, 6 lateral and 2 dorsal tactiles, 2 blunt sensory setae, dt 1, 2
solid, nude, anterior dt 3 solid, nude (hollow, nude), posterior dt 3
hollow, nude; tarsus III, 12 (11) ventrals, 6 laterals, trichoboth,
dorsoterminals as in leg II; tarsus IV, 14 (12-14) ventrals, 7 (6)
laterals, trichoboth, dt 1, 2 solid, nude, dt 3 hollow, nude.

Male: Unknown.

Type: Tritonymph, Planada, California, June 13, 1936, E. W.
Baker, lichens from fig tree.

Location of type: The United States National Museum, type no.
1462.

Remarks: One hundred and forty-six females were examined,
which included specimens from: Mexico (Mexico, Tamaulipas,
Guerrero, Neuvo Leon, San Luis Potosi) and the United States
(Texas, California, Utah, Colorado, Washington, Alabama, Mary-
land).

This is the only species studied in which a marked variation oc-
curs in the pattern of the dorsal propodosomal shield. The striae
in the central area of the pattern in the Mexican specimens are
oblique (fig. 159), while the same striae in the United States (Utah)
specimens are longitudinally directed (fig. 160). Variation in the
forms of the dorsoterminal setae at the apex of the tarsi can not
be related to geographic area, nor does correlation exist between
the different striation patterns and the various combinations of
dorsoterminal setae.

The redescription and figures are based on a female with the
following data: Cedar Mountains, Tooele County, Utah, Decem-
ber 12, 1954, D. Allred, Ex: pack rat (Neotoma) nest. The speci-
men used for the illustration of the Mexican shield pattern was
collected at: 6 miles northeast of Jalostitlan, Jalisco, Mexico,
August 19, 1954, W. T. Atyeo, under rock, altitude 6,000 feet.

434 The University Science Bulletin

Spinibdella ornata, sp. nov.

(Figs. 64, 121, 122, 161, 176)
This species is closely related to Spinibdella hifurcata, sp. nov.,
but is unique in having the duplex setae on the proximal halves of
the genua, rather than the distal halves, and one blunt and one
attenuate sensory seta on tibia II, rather than one deeply recessed
blunt sensory seta. The internal humerals of Spinibdella ornata
are about one half the length of the first interspaces, whereas in
S. bifiircata, the internal humerals equal the first interspaces. Males
can be easily recognized by the branched setae on the amphioid
sclerites (fig. 176).

Female: Color unknown. Body narrow, strongly constricted;
length, including gnathosoma, 900 \j.. Gnathosoma: Length, 252 [a;
palpus (fig. 64) short; measurements: I, 10 \j.; II plus III, 167 jj.;
IV, 21 [x; V, 36 [x; des, 245 [jl; ves, 167 [j.. Chelicera striated, 239 [x in
length; setae as in fig. 33. Gnathosoma striated; dorsal hypostomal
setae, 25 \^ in length. Dorsal propodosoma (fig. 161): Striae
sparsely broken; lateral propodosomals nude, 41 pi in length; eyes
separated by distance equal to diameter of anterior pair, interval
between eyes with longitudinal striae; distance between anterior
sensilla, 53 \i. Dorsal hysterosoma: Setae finely branched; length of
internal humeral, 37[ji., approximately one half of first interspace;
external humeral, 47 \i in length; sacrals and clunals in subtrapa-
zoidal arrangement. Anal region: Border striae parallel, 2 pairs
of anal setae, paranals wanting, postanals flanking termination of
cleft. Genital region: Each genital plate with 10 small, attenuate
setae in linear arrangement; 18 pairs of paragenitals, anterior pair
between coxae IV; 2 median, unpaired setae anterior to genital
aperture; genital discs equidistant, in posterior two thirds of
vestibule; ovipositor with 12 subapical, 6 postmedial setae. Legs
(figs. 121, 122): Claws with one row of lateral rays each; measure-
ments: tibia I, 72 [x; tarsus I, 96 \i; tibia II, 61 [x; tarsus II, 90 [a.
Chaetotaxy : coxae I-IV, 8, 7, 7, 6 tactile setae, length of each seta
exceeding coxal width; trochanters I-IV, 1, 1, 2, 1 tactile setae;
basifemora I-IV, 12, 8, 7, 3 tactile setae; telofemora I-IV, 9, 8, 5, 6
tactiles; genu I, 6 tactiles, 1 proximal duplex seta, 1 distal attenuate
sensory seta; genua II-III, 6 tactiles and 1 proximal duplex seta
each; genu IV, 8 tactiles; tibia I, 15 tactiles, 1 attenuate and 1
blunt sensory seta, 1 peg, trichoboth; tibia II, 13 tactiles, 1 attenuate
and 1 blunt sensory seta; tibia III, 12 tactiles, 1 attenuate sensory
seta; tibia IV, 15 tactiles, trichoboth; tarsus I, 17 ventrals, 8 lateral

A Revision of the Mite Family Bdellidae 435

and 2 dorsal tactile setae, 1 attenuate and 2 blunt sensory setae, 1
peg, dt 1, 3 hollow, nude, dt 2 solid, nude; tarsus II, 17 ventrals, 7
lateral and 1 dorsal tactile seta, 1 attenuate and 2 blunt sensory setae,
1 peg, dt 1, 2 solid, nude, dt 3 hollow, nude; tarsus III, 16 ventrals,
6 laterals, trichoboth, dorsoterminals as in tarsus II; tarsus IV, 17
ventrals, 5 laterals, trichoboth, setae of dt 1 obliquely situated to
each other, dt 2, 3 as in tarsus II.

Male: Identical to female except in genital region: each genital
plate with 17 equal setae, each about one half the length of female
genital setae; amphioid sclerites as in fig. 176.

Holotype: Male, Bear Lake, Rocky Mountain National Park,
Colorado, August 23, 1955, T. A. Woolley, moss and litter.

Allotype: Female, same data as holotype.

Paratypes: One female, same data as holotype; one female, Hope
Valley, Alpine Co., California, July 22, 1955, N. A. Walker, lodge
pole pine litter and humus, altitude 7,300 feet; one male, H. Cowell,
Redwood State Park, 4 miles north of Santa Cruz, Santa Cruz Co.,
California, August 31, 1956, N. A. Walker, redwood litter, sample
294; one female, 3 miles north of Boulder Creek, Santa Cruz Co.,
California, August 31, 1956, N. A. Walker, redwood litter, sample
295.

Location of types: The holotype and three paratypes are de-
posited in the Snow Entomological Museum; the allotvqje at the
Department of Zoology, Colorado A and M College, Fort Collins,
Colorado; and one paratype is deposited at the United States Na-
tional Museum.

Remarks: Variation between the Colorado and California speci-
mens is not detectable. Total length varies from 660 \i to 923 [ji,
and the other measurements vary accordingly. This species is
named ornata to call attention to the highly modified setae on the
male amphioid sclerites. Drawings of the holotype.

Genus Biscirtis Thor

Biscims Thor, 1927, Ann. Mus. Leningrad, vol. 27, p. 135. (Type: Bdella
silvatica Kramer, 1881 [by original designation!.)

This genus is related to Monotrichohdclla, but has two end setae
rather than one. The palpal tibiotarsus is elongated and cylindri-
cal, and in addition to the two end setae, has two other setae, as
does the genu and the basifemur. The cheliceral setae do not ex-
tend beyond the tips of the chelae. The lateral propodosomal

436 The University Science Bulletin

setae are wanting; striation patterns on the dorsal propodosoma
do not appear to be specific.

The majority of the species formally placed in this genu (Thor,
1931) appear to belong to Thoribdella; this would include such
species as Biscirus lapidarius (Kramer), 1881; B. intermedins Thor,
1928; and B. imcinatiis (Kramer), 1881. Only two species have
been examined in the present study that definitely belong to this
group, these are: Biscirus silvaticus (Kramer), 1881, and B. thori
Womersley, 1933.

Biscirus silvaticus (Kramer)
(Figs. 10, 35, 67, 129, 130, 166, 177)
Bdella silvatica Kramer, 1881, Zeitsch. fiir Naturw., vol. 54, p. 445.

This species is related to the Australian species, Biscirus thori
Womersley, but can be distinguished by the palpal genu being
less than twice the length of the palpal telofemur, rather than the
genu three times longer than the telofemur.

Female: Color in hfe dark red with deep blue blotches and black
to purple eye spots. Body strongly constricted; length, including
gnathosoma, 1210 \j. (1100-1800 \i). Gnathosoma: Length, 349 \i
(349-485 \i); palpus (fig. 67) with reduced number of setae; meas-
urements: I, 14 ix (11-17 [x); 11,175 !x (175-491 [x); III, 29 |x (26-
54 [x); IV, 43 [X (43-95 ix); V, 126 ix (114-221 [x); des, 235 [x (235-
280 [x); ves, 170 iJ. (170-207 ix). Chehcera (fig. 10) elongate,
striated, 345 [x (331-460 [x) in length; setae as figured. Hypostome
striated, 349 [). ( 349-485 [). ) in length; dorsal hypostomal setae want-
ing. Dorsal propodosoma (fig. 166): Striae coarsely broken;
median propodosomals plumose, 65 [x (44-65 [x) in length; eyes sep-
arated by distance equal to diameter of anterior pair, interval be-
tween eyes with transverse striae; distance between anterior sen-
silla, 65 [X (65-103 \).). Dorsal hysterosoma: Setae plumose; length
of internal humeral, 51 [x (37-51 (x), approximately one third of
first interspace; external humeral, 61 [x (37-61 [x) in length; sacrals
and clunals in gently curving transverse rows. Anal region: Bor-
der striae parallel; one pair of anal setae; 2 pairs of paranals;
postanals flanking termination of cleft, equal in length to external
clunals. Genital region: Each genital plate with 15 (14) thin, nude
genital setae arranged in 2 irregular rows; 12 (13) pairs of para-
genitals, anterior pair between coxae IV; genital discs of moderate
size, equidistant; ovipositor with 14 subapical setae. Legs (figs.

A Revision of the Mite Family Bdellidae 437

129, 130): Claws shorter than pretarsus, each with one row of
lateral rays; measurements: tibia L 102 \). (102-158 \i); tarsus I,
177 PL (170-218 (x); tibia II, 96 [i (88-129 \i); tarsus II, 170 \i (160-
207 ii). Chaetotaxy: coxae I-IV, 4, 4, 5, 4 tactile setae; trochan-
ters I-IV, 1 tactile seta each; basifemora I-IV, 8, 9, 7, 4 tactile
setae; telofemora I-IV, 5, 5, 4, 3 tactile setae; genu I, 3 tactiles,
4 attenuate sensory setae; genu II, 4 tactiles, 1 attenuate sensory
seta; genu III, 4 tactile setae, 1 attenuate sensory seta; genu IV,
4 tactile setae; tibia I, 12 tactile setae, 4 attenuate sensory setae,
1 peg, trichoboth; tibia II, 13 tactile setae, 1 attenuate and 1 blunt
sensory seta; tibia IV, 11 tactile setae, trichoboth; tarsus I, 15
plumose ventrals, 6 lateral and 1 dorsal tactile seta, 2 attenuate
and 2 blunt sensory setae, dt 1, S hollow, nude, anterior dt 2 want-
ing, posterior dt 2 solid, plumose; tarsus II, 14 plumose ventrals,
6 lateral and 2 dorsal tactiles, 2 blunt sensory setae, dt 1, 2 solid,
nude, dt 3 hollow, nude; tarsus III, 16 plumose ventrals, 18 laterals,
trichoboth, dorsoterminals as in tarsus II; tarsus IV, 13 plumose
ventrals, 7 laterals, trichoboth, 1 attenuate sensory seta proximal
to trichoboth, dt 1 reduced to one solid, nude seta, dt 2, 3 as in
tarsus II.

Male: Identical to female except in genital region; each genital
plate with 19-20 setae; amphioid sclerites as in fig. 177.

Type: Europe.

Location of type: Unknown.

Remarks: Specimens were examined from: Mexico (Oaxaca,
Durango, Nuevo Leon), Haiti, the United States (California, Mis-
sissippi, Kansas, Colorado, Maryland, Tennessee), Iceland, and
Germany (near Ost-Holstein ) .

Typically, the median propodosomals divide the interval between
the posterior sensilla into three approximately equal parts, but con-
siderable migration occurs around these points. The median pro-
podosomals can be anterior to a line connecting the posterior sensilla
(fig. 166) or they can be on that line, and can be closer to the pos-
terior sensilla than to themselves, or vice versa.

Redescription and figures of a female from: ten miles south of
China, Nuevo Leon, Mexico, August 4, 1955, R. E. Beer, under rock.
The figure of the amphioid sclerites was prepared from a male from :
University of Kansas campus, Lawrence, Douglas Co., Kansas, May
5, 1954, D. E. Sbur, under rock.

438 The University Science Bulletin

Genus Monotrichobdella Baker and Balock

Monotrichobdella Baker and Balock, 1944, Proc. Ent. Soc. Washington, vol.
46, no. 7, p. 176. (Type: Monotrichobdella max-osburni Baker and Balock,
1944 [by original designation].)

This unique genus is related to Biscirus, but has one palpal end
seta rather than two. The palpal tibiotarsus is elongate and cylin-
drical, and in addition to the single apical seta, there are three other
setae on this segment; the genu may have one or two setae and the
basifemur has two setae. The distal cheliceral seta extends beyond
the tips of the chela. The lateral propodosomal setae are wanting;
striation patterns on the dorsal propodosoma are similar to Biscirus
species.

Monotrichobdella maxosburni Baker and Balock

(Figs. 34, 68, 131, 132, 168)

Monotrichobdella max-osburni Baker and Balock, 1944, Proc. Ent. Soc. Wash-
ington, vol. 46, no. 7, p. 176.

To date this is a monotypic genus. Adults have not been ex-
amined, but nymphal forms have been studied from two localities.

Tritonymph: Color in hfe red. Body ovoid, weakly constricted;
length, including gnathosoma, 1350 \i. Gnathosoma: Length, 341 [jl;
palpus (fig. 68) with distal two thirds of tibiotarsus extending be-
yond hypostome; measurements: I, 11 [;.; II plus III, 211 pi; IV, 31 [jl;
V, 102 [jl; end seta, 216 ]s.. Chehcera (fig. 34) elongate, finely stri-
ated, 210 ^ in length; chela smooth, needlelike, fixed digit two thirds
length of movable digit; setae as figured. Gnathosomal base
striated; hypostome nonstriated; vh i, 65 pi from apex; vh 2, 218 [x
from apex; dorsal hypostomal setae wanting. Dorsal propodosoma
(fig. 168): Coarsley broken striae; median propodosomals plumose,
54 [A in length; eyes separated by distance equal to diameter of the
anterior pair; distance between anterior sensilla, 82 pi. Dorsal
hysterosoma: Setae plumose; length of internal humeral, 58 [x, ap-
proximately one half of first interspace; external humeral, 58 pi in
length; sacrals in strongly curving transverse row; external clunals
wanting. Genital region: Each genital plate with 9 fine, equal setae
in irregular linear arrangement; 11 pairs of paragenitals, 1 unpaired
seta and anterior 2 pairs of paragenitals between coxae IV; genital
discs small, anterior pair one third the length of vestibule from
anterior end, posterior pairs in caudal third; genitalia wanting.
Anal region: Border striae parallel; 2 pairs of anal setae; 2 pairs
of paranals; postanals flanking termination of cleft, shorter than
clunal setae. Legs (figs. 131, 132): Each claw with one row of

A Revision of the Mite Family Bdellidae 439

lateral rays; measurements: tibia I, 85 [x; tarsus I, 134 \l; tibia II,
82 [i; tarsus II, 128 \j.. Chaetotaxy: coxae I-IV, 4, 3, 4, 3 tactile
setae; trochanters I-IV, 1 tactile seta each; basifemora I-IV, 7, 7,
7, 4 tactile setae; telofemora I-IV, 5, 5, 4, 4 tactile setae; genua
I-III, 4 tactile setae and 1 attenuate sensory seta each; genu IV,
4 tactile setae; tibia I, 8 tactile setae, 3 attenuate sensory setae, 1
solid, attenuate peg, trichoboth; tibia II, 9 tactile setae, 1 attenuate
sensory seta, 1 recessed, blunt sensory seta; tibia III, 8 tactile
setae, 1 attenuate sensory seta; tibia IV, 6 tactile setae, trichoboth;
tarsus I, 10 plumose ventral setae, 6 lateral and 1 dorsal tactile
seta, 2 attenuate and 2 blunt sensory setae, 1 solid, attenuate peg,
dt 1, 2 solid, nude, dt 3 hollow, nude; tarsus II, 10 plumose ven-
trals, 6 lateral and 1 dorsal tactile seta, 2 blunt sensory setae,
dorsoterminals as in tarsus I; tarsus III, 10 plumose ventrals, 6
laterals, trichoboth, dorsoterminals as in tarsus I; tarsus IV, 10
plumose ventrals, 4 laterals, trichoboth, 1 attenuate sensory seta
proximal to trichoboth, dorsoterminals as in tarsus I.
Female, Male: Not examined.

Holofype: Tritonymph, near Tres Cumbres, Mexico-Cuernavaca
Highway, Morelos, Mexico (alt. 10,000 ft.), January 7, 1943, J. W.
Balock and J. G. Shaw, lichens.

Location of type: The United States National Museum, type no.
1459.

Remarks: In addition to the type, two other specimens were
available: one deutonymph, same data as the holotype, and one
protonymph from: Salazar, Distrito Federal, Mexico, F. Bonet,
September 30, 1942, moss.

The unpaired paragenital seta is not present in the protonymphal
and deutonymphal stages. In other species having this unpaired
seta, it is present in all stages of .development, therefore, it is pos-
sible that the holotype is an aberrant specimen and that a median,
unpaired paragenital seta is not a characteristic of this species, or
that the unpaired seta occurs in only the tritonymphal and adult
stages. Drawings of the holotype.

SELECTED BIBLIOGRAPHY
Andre, M.

1947. Croisiere du Bouganville aux iles Australes francaises-XMI Acaricns.
Mem. Mus. national hist, nat., n. s., vol. 20, p. 89.
Atyeo, Warren T.

1958. The genus Bonzia in the new world (Acarina, Cunaxidae). J. Kan-
sas Ent. Soc., vol. 31, no. 2, pp. 173-177.

440 The University Science Bulletin

Baker, E. W., and J. Balock.

1944. Mites of the family Bdellidae. Proc. Ent. Soc. Washington, vol.
46, no. 7, pp. 176-184.
Beer, Robert E.

1954. A revision of the Tarsonemidae of the Western Hemisphere ( Order,
Acarina). Univ. Kansas Sci. Bull., vol. 36, part 2, no. 16, pp.
1109-1112.

BOTTAZZI, E.

1950, Primo contributo alia fauna de Trombidiformes (Acari) del Par-
mense, Monit. Zool. ital., Firenze, vol. 58, no. 1-6, p. 31.
Cooreman, J.

1943. Note sur la faune des Hautes-Fagnes en Belgique. XII. Acariens
(Trombidiformes). Bull. Mus. Hist. Nat. Belgique, vol. 19, no.
64, p. 7.

CtTRREE, G. A.

1934. The Bdellid mite Biscirus lapidarius Kramer, predatory on the
lucerne flea Sminthurus viridis L. in Western Australia. J. Aus-
tral. Council Sci. & Indus. Res., vol. 7, no. 1, pp. 9-20.
EWTNG, H. E.

1937. Insects and arachnids from Canadian Amber. Univ. Toronto Stud.,
Geol. Ser. no. 40, pp. 57-60.

Evans, G. Owen

1952. Terrestrial Acari new to Britain. II. Ann. Mag. Nat. Hist., ser.
12, vol. 5, p. 668.

1953. On a collection of Acari from Kilimanjaro (Tanganyika). Ann.
Mag. Nat. Hist., ser. 12, vol. 6, pp. 272-274.

Garman, Philip

1948. Mite species from apple trees in Connecticut. Connecticut Ag.
Exper. Sta., New Haven, Bull. 520, pp. 22-24.
Geoffroy, Etienne Louis

1762. Histoire abregee des Insects, qui se trouvent aux Environs de
Paris; dans laquelle ces Animaux sont ranges suivant un Ordre
methodique, Paris, vol. 2, p. 617.
Grandjean, F.

1938. Observations sur les Bdelles (Acariens). Ann. Soc. Ent. France,
vol. 107, pp. 1-24.

Hermann, Jean-Frederic

1804. Memoire apterologique. Ouvrage couronne en 1790 par la Societe

d'Histoire naturelle de Paris, PubUe par Frederic-Louis Hammer,

Strasbourg, pp. 60-62.
Irk, V.

1939. Die terricolen Acari der Otztaler um Stubaier Hochalpen. Veroff.
Mus. Ferdinand, Innsbruck, vol. 19, p. 158.

Jacot, a. p.

1938. Thomas Say's free-hving mites re-discovered. Psyche, vol. 45, pp.
126-130.
Latreille, Pierre Andre

1795. Observations sur la variete des organes de la bouche des Tiques, et
distribution methodique des insectes de cette famille d'apres les

A Revision of the Mite Family Bdellidae 441

caracteres establis sur la conformation de ces organs. Magasin
encyclopedique, ou Journal des Sciences, des Lettres et des Arts,
redige par Millin, Noel et Warens, Paris, vol. 4, p. 18.
1796. Precis des Caracteres generiques des Insects, diposes dans un
Ordre naturel, Paris, p. 180.
LiPovsKY, Louis J.

1953. Improved technique for rearing chigger mites (Acarina: Trom-
biculidae). Ent. News, vol. 64, pp. 4-7.
McGregor, E. A.

1956. The mites of citrus trees in southern California. Mem. Southern
California Acad. Sci., vol. 3, no. 3, pp. 12-13.
Michael, A. D.

1896. The internal anatomy of Bdella. Trans. Linn. Soc. London, ser,
2, vol. 6, Zool, pp. 477-528.

OUDEMANS, A. C.

1937. Kritisch Historisch Overzicht der Acarologie. E. J. Brill, Leiden,
vol. 3, part C, pp. 1179-1248.
Pritchard, a. E., and Baker, E. W.

1955. A revision of the spider mite family Tetranychidae. Pacific Coast
Ent. Soc, Mem. ser., vol. 2, pp. 5-11.

SCHWEIZER, J.

1951. Die Landmilben des Schweizerischen Nationalparkes. 2 Teil:
Trombidiformes Renter 1909. Ergben. wiss. Unters. schweiz.
Nationalparkes N. F., vol. 3, no. 23, p. 74.
Snetsinger, Robert

1956. Biology of Bdella depressa, a predaceous mite. J. Ec. Ent., vol.
49, no. 6, pp. 1^5-1 AQ.

Thor, Sig

1928. Norwegische Bdellidae III und einige Bemerkungen iiber Haare,

Chitinleisten und Schildcr am Cephalothorax. Zool. Anz., vol. 77,

no. 9/10, pp. 213-219.
1931a. Bdellidae, Nicoletiellidae, Cryptognathidae. Das Tierreich, vol.

56, pp. 1-64.
1931b. Nordafrikanische Bdellidae und Cunaxidae von Dr. F. Grandjean

(Paris) gesammelt. Zool. Anz., vol. 97, no. 1/2, pp. 62-76.
1937. Iloploscirus, eine neue schildtragende Bdellidcngattung aus Tas-
mania. Zool. Anz., vol. 119", no. 1/2, pp. 43-44.
Tragardh, Ivar.

1902. Littoralen Arten der Gattung Bdella Latr. Bihang Till K. Svenska

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ViTZTHiTM, H. Graf

1931. Resultats Scientifiques du Voyage aux Indes Orientales Neer-

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442 The University Science Bulletin

1933. Terrestrische Acarienen aus Mexiko. Zool. Anz., vol. 103, no.

9/10, pp. 226-228.
1940-1943. Acarina. Bronns' Klassen und Ordungen des Tierreiches 5,
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WlLLMANN, C.

1939a. Die Arthropodenfauna von Madeira. XIV. Terrestrische Acari

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Ver. Bremen, vol. 31, no. 3, p. 532.
1941. Die Acari der Hohlen der Balkanhalbinsel (Nach dem Material

der "Biospeologica balcanica"). Stud. Geibiete Allg. Karstf.

Berlin Biol. Ser., no. 8, p. 58.
1951. Die hochalpine Milbenfauna der mittleren Hohen Tauem insbe-

sondere des Grossglockner-Geietes (Acari). Bonn. Zool. Beitr.,

vol. 2, p. 162.

WOMERSLEY, H.

1933a. A preliminary account of the Bdelhdae (Snout Mites) of Aus-
tralia. Trans. Roy. Soc. South Australia, vol. 57, pp. 97-107.

1933b. On some Acarina from Australia and South Africa. Trans. Roy.
Soc. South Australia, vol. 57, p. 111.

1933c. A possible biological control of the clover springtail or lucerne flea
{Sminthurus viridis L. ) in Western Australia. J. Austral. Council
Sci. & Indus. Res., vol. 6, no. 2, pp. 83-91.

444 The University Science Bulletin

Figure 1

Fig. 1. Dorsal aspect of SpinibdeUa bifurcata, sp. nov.

a. chelicera

b. gnathosomal base

c. anterior sensillum

d. lateral propodosomal

e. lateral eye

f. median propodosomal

g. posterior sensillum
h. internal humeral
i. external humeral
j. internal dorsal
k. internal lumbral
1. internal sacral
m. external sacral

n. internal clunal (

o. posterior anal i

p. anal seta 'jl

q. anal cleft I

A Revision of the Mite Family Bdellidae

445

Figure 1

446

The University Science Bulletin

Figure 2

Fig. 2. Ventral aspect of Spinibdella bifurcata, sp. nnv.

a. adornal setae

b. lateral lip

c. ventral hypostomal seta

d. buccal cone

e. gnathosomal base

f. coxal peg

g. paragenital seta
h. genital plate

1. genital seta
j. anal seta

A Revision of the Mite Family Bdellidae

447

Figure 2

448 The University Science Bulletin

Figures 3-11 i

Musculature of the appendages (figs. 3-6) ||

Fig. 3. Leg of Bdella species
Fig, 4. Leg of Neomolgus species
Fig. 5. Palpus of Bdella species
Fig. 6. Palpus of Cunaxa species

Legends for Figures 3-6

a. flexor muscle of genu [

b. flexor muscle of tibia '

c. flexor muscle of tarsus |

d. depressor muscle of pretarsus

e. elevator muscle of pretarsus I

f. extensor muscle of genu .
C. coxa

Tr. trochanter j

BF. basifemur ■'

TF. telofemur

G. genu '

Tb. tibia

Ta. tarsus |

TbTa. fused tibiotarsus ,

Fig. 7. Chela of Bdella longicomis \

a. fixed digit

b. movable digit

Fig. 8. Chela of Thoribdella simplex

Fig. 9. Chela of Odontoscirus iota

Fig. 10. Chela of Biscirtis silvaticus {

Fig. IL Chela of Cyta spuria '

A Revision of the Mite Family Bdellidae
Figures 3-11

449

15—8920

450 The University Science Bulletin

Figures 12-23

Lateral aspects of right chelicerae

Fig. 12. Bdella longicornis

Fig. 13. Bdellu mexicana

Fig. 14. Bdellodes longirostris

Fig. 15. Odantoscirus alpinus, holotype

Fig. 16. Odontoscirus iota, holotype

Fig. 17. Odontoscirus iota, holotype

Fig. 18. Octobdellodes hurdi, holotype

Fig. 19. Octobdellodes infrequens, holotype f

Fig. 20. Octobdellodes infrequens, allotype ,

Fig. 21. Tharibdella meridionalis

Fig. 22. Thoribdella communis, holotype

Fig. 23. Thoribdella simplex, holotype

A Revision of the Mite Family Bdellidae
Figures 12-23

451

too^

452 The University Science Bulletin

Fig.

24.

Fig.

25.

Fig.

26.

Fig.

27.

Fig.

28.

Fig.

29.

Fig.

30.

Fig.

31.

Fig.

32.

Fig.

33.

Fig.

34.

Fig.

35.

Figures 24-35

Lateral aspects of right chelicerae

Thoribdella truncata, holotype
Thoribdella califomica
Thoribdellu insolita, holotype
Thoribdella spinosa, holotype
Neomolgus mutabilis, holotype
Neomolgus littoralis
Cyta latirostris
Cyta coerulipes
Spinibdella tenuirostris, type
Spinibdella corticis
Monotrichobdella maxosburni, type
Biscirus silvaticus

A Revision of the Mite Family Bdellidae
Figures 24-35

453

to«^

454

The University Science Bulletin

Figures 36-41

Median aspects of left palpi

Fig.

36.

Bdella mexicana, type

Fig.

37.

Bdella longicornis

Fig.

38.

Bdella longistriata, holotype

Fig.

39.

Bdella muscorum

Fig.

40.

Bdella tropica, holotype

Fig.

41.

Bdella distincta, lectotype

A Revision of the Mite Family Bdellidae
Figures 36-41

455

456 The University Science Bulletin

Figures 42-47 ]

Median aspects of left palpi j

Fig. 42. Bdellodes longirostris j

Fig. 43. Odontoscirus alpinus, holotype 1

Fig. 44. Odontoscirus iota, holotype J

Fig. 45. Octohdellodes hurdi, holotype j

Fig. 46. Octohdellodes infrequens, holotype I

Fig. 47. Bdellodes bisetosa, holotype i

A Revision of the Mite Family Bdellidae

457

Figures 42-47

458 The University Science Bulletin

FiGLTRES 48-54

Median aspects of left palpi

Fig. 48. Thoribdella communis, holotype

Fig. 49. Thoribdella simplex, holotype

Fig. 50. Thoribdella truncata, holotype

Fig. 51. Thoribdella meridionalis

Fig. 52. Thoribdella californica

Fig. 53. Thoribdella insolita, holotype

Fig. 54. Thoribdella spinosa, holotype

A Revision of the Mite Family Bdellidae

459

Figures 48-54

460 The University Science Bulletin

Figures 55-60

Median aspects of left palpi
Fig. 55. Neomolgus mutabilis, holotype
Fig. 56. Neomolgus littoralis
Fig. 57. Cyta coerulipes, nude setae
Fig. 58. Cyta coerulipes, plumose setae
Fig. 59. Ctjta spuria, holotype
Fig. 60. Cyta latirostris

A Revision of the Mite Family Bdellidae

461

Figures 55-60

462 The University Science Bulletin

Figures 61-68

Median aspects of left palpi
(Figs. 61-65, scale A; figs. 66-68, scale B)
Fig. 61. SpinibdeUa tetiuirostris, type
Fig. 62. SpinibdeUa depressa
Fig. 63. SpinibdeUa bifurcata, holotype
Fig. 64. SpinibdeUa ornata, holotype
Fig. 65. SpinibdeUa cronini
Fig. 66. SpinibdeUa coiticis
Fig. 67. Biscirus silvaticus
Fig. 68. MonotrichobdeUa maxosburni, type (tritonymph)

A Revision of the Mite Family Bdellidae
Figures 61-68

463

464 The University Science Bulletin

Figures 69-72
Dorsal aspect of legs I and II of Bdella longicomis
Fig. 69. Distal tliree segments of leg I

Fig. 70. Enlarged claw illustrating large and small lateral rays
Fig. 71. Tarsus I, stippled areas indicate extent of setal migration
Fig. 72. Distal three segments of leg II

A Revision of the Mite Family Bdellidae

Figures 69-72
70

465

466 The University Science Bulletin

Figures 73-78
Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
Figs. 73, 74. Bdella muscorum
Figs. 75, 76. Bdella longistriata, holotype
Figs. 77, 78. Bdella tropica, holotype

A Revision of the Mite Family Bdellidae

467

Figures 73-78

lOO^

468 The University Science Bulletin

Figures 79-84

Dorsal aspect of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
(Figs. 79-82, scale A; figs. 83-84, scale B)
Figs. 79, 80. Bdella distincta, lectotype
Figs. 81, 82. Bdella mexicana, type
Figs. 83, 84. Odontoscirus iota, holotype

A Revision of the Mite Family Bdellidae 469

Figures 79-84

lOOp 200^

A •

470 The University Science Bulletin

Figures 85-90

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
Figs. 85, 86. Odontoscirus alpinus, holotype
Figs. 87, 88. Bdellodes longirostris
Figs. 89, 90. Octohdellodes infrequens, holotype

A Revision of the Mite Family Bdellidae
Figures 85-90

471

-ZOOii

472 The University Science Bulletin

Figures 91-96 1

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)

Figs. 91, 92. Bdellodes bisetosa, holotype \

Figs. 93, 94. Octobdellodes htirdi, holotype j

Figs. 95, 96. Thoribdella meridionalis '

A Revision of the Mite Family Bdellidae
Figures 91-96

473

.^^^^.^^

— 200/1

^^^4 The University Science Bulletin

Figures 97-102
Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
Figs. 97, 98. Thoribdella communis, holotype
Figs. 99, 100. Thoribdella insolita, holotype
Figs. 101,102. Thoribdella trtincata, holotype

A Revision of the Mite Family Bdellidae

475

Figures 97-102

20C|l

476 The University Science Bulletin

Figures 103-108

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
Figs. 103, 104. Thoribdella simplex; leg I, Kansas paratype; leg II, holotype
Figs. 105, 106. Thoribdella califomica
Figs. 107, 108. Neomolgus littoralis

A Revision of the Mite Family Bdellidae

477

Figures 103-108

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-200|i

478 The University Science Bulletin

Figures 109-114
Dorsal aspects of the three distal segments of legs I and 11
(Legs I, odd numbers; legs II, even numbers)
Figs. 109, 110. Thoribdella spinosa, holotype
Figs. Ill, 112. Neomolgus mutabilis, holotype
Figs. 113, 114. Cyta spuria, holotype

A Revision of the Mite Family Bdellidae

479

Figures 109-114

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I "si^ '^

200^

480 The University Science Bulletin

Figures 115-120

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
(Figs 115, 116, scale A; figs. 117-120, scale B)
Figs. 115, 116. Cijta coerulipes
Figs. 117, 118. Spinibdella bif areata, holotype
Figs. 119, 120. Spinibdella depressa

A Revision of the Mite Family Bdellidae
Figures 115-120

481

A •-

B»-

16—8920

20 0/1
lOO/i

482 The University Science Bulletin

FiGUBES 121-126

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even numbers)
Figs. 121, 122. Spinibdella ornata, holotype
Figs. 123, 124. Spinibdella tenuirostris, type
Figs. 125, 126. Spinibdella cronini

A Revision of the Mite Family Bdellldae 483

Figures 121-126

100)1

484 The University Science Bulletin

Figures 127-132

Dorsal aspects of the three distal segments of legs I and II
(Legs I, odd numbers; legs II, even nmnbers)
(Figs. 129-130, scale A; figs. 127-128, 131-132, scale B)
Figs. 127, 128. Spinibdella coHicis
Figs. 129, 130. Biscinis silvaticus
Figs. 131,132. Monotrichobdella 7naxosburni, type (tritonymph)

A Revision of the Mite Family Bdellidae

485

Figures 127-132

486 The University Science Bulletin

Figures 133-141
Striation patterns on the dorsum of the propodosoma
Fig. 133. Bdella longicarnis from Cortez Pass, Mexico

a. anterior pseudostigmatic organ

b. lateral internal apodeme

c. insertion of the lateral propodosomal seta

d. minor internal apodemes

e. posterior pseudostigmatic organ

f. insertion of the median propodosomal seta
Fig. 134. Bdella longicomis from Cortez Pass, Mexico
Fig. 135. Bdella longicomis from Cortez Pass, Mexico
Fig. 136. Bdella muscorum from Douglas Lake, Michigan
Fig. 137. Bdella muscorum from N. W. Arkansas

Fig. 138. Bdella muscorum from Douglas Lake, Michigan
Fig. 139. Bdella muscorum from Iceland
Fig. 140. Bdella tropica, holotype
Fig. 141. Bdella distincta, lectotvpe

A Revision of the Mite Family Bdellldae

487

Figures 133-141

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140

aoqp

488 The University Science Bulletin

^,1

Figures 142-150

Modifications of the posterior sensilla and median propodosomal setae of

Thoribdella species

General striation pattern of Thoribdella species

Thoribdella spinosa, holotype

Thoribdella californica

Thoribdella meridionalis

Thoribdella communis, holotype

Thoribdella simplex, holotype

Thoribdella insolita, holotype

Thoribdella truncata, holotype

Thoribdella truncata, enlargement of posterior sensillum

Fig.

142.

Fig.

143.

Fig.

144.

Fig.

145.

Fig.

146.

Fig.

147.

Fig.

148.

Fig.

149.

Fig.

150.

A Revision of the Mite Family Bdellidae

489

Figures 142-150

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142

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490 The Unwersity Science Bulletin

Figures 151-156

Striation patterns on the dorsum of the propodosf^ma

Fig. 151. Bdella longistriata, holotype ]

Fig. 152. Octobdellodes hurdi, holotype 1

Fig. 153. Octobdellodes infrequens, holotype )

Fig. 154. Odontoscirus iota, holotype j

Fig. 155. Bdellodes bisetosa, holotype j

Fig. 156. Spinibdella corticis '\

A Revision of the Mite Family Bdellidae

491

Figures 151-156

\ >,v^ N w^^ o - ~ - - r ''

154

153

20 0>

156

492 The University Science Bulletin

Figures 157-162
Striation patterns on the dorsum of the propodosoma
Fig. 157. Spinibdella tenuirostris, type
Fig. 158. Spinibdella depressa
Fig. 159. Spinibdella cronini from Mexico
Fig. 160. Spinibdella cronini from Utah
Fig. 161. Spinibdella omata, holotype
Fig. 162. Spinibdella bifurcata, holotype

A Revision of the Mite Family Bdelledae

493

Figures 157-162

lis

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157

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158

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160

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161

20<!ti

494 The University Science Bulletin

Figures 163-168

Striation patterns on the dorsum of the propodosoma

Fig. 163. Neomolgus mutabilis, holotype

Fig. 164. Cyta latirostris

Fig. 165. Cijta coerulipes

Fig. 166. Biscirus silvaticus

Fig. 167. Cyta spuria, holotype

Fig. 168. MonotrichobdeUa maxosburni, type (tritonymph)

A Revision of the Mite Family Bdellidae

495

Figures 163-168

164

163

,i{/^fc#^®

^5^^'

166

^im

167

496 The University Science Bulletin

Figures 169-177

Dorsal aspects of the right amphioid sclerites

Fig.

169.

Bdella longicornis

Fig.

170.

Bdella longistriata, holotype

Fig.

171.

Ctjta coerulipes

Fig.

172.

Cyta latirostris

Fig.

173.

Cyta spuria, holotype

Fig.

174.

Spinibdella bifurcata, holotype

Fig.

175.

Spinibdella depressa

Fig.

176.

Spinibdella ornata, holotype

Fig.

177.

Biscirus silvaticus

A Revision of the Mite Family Bdellidae

497

Figures 169-177

498 The University Science Bulletin

Figures 178-197

Fig. 178. Genital region of Cyta latirostris

a. genital seta

b. paragenital seta

c. unpaired median paragenital seta

d. genital trachea

e. genital disc

f. genital spine

Types of setae found in the Bdellidae

Fig. 179. Tactile seta, solid, pilose

Fig. ISO. Tactile seta, solid, coarsely pilose

Fig. 181. Tactile seta, soHd, branched

Fig. 182. Tactile seta, solid, branched distally

Fig. 183. Tactile seta, solid, plumose

Fig. 184. Trichoboth ( long sensory seta of the legs )

Fig. 185. Dorsoterminal seta of legs, hollow, nude

Fig. 186. Dorsoterminal seta of legs, hollow, minutely pilose

Fig. 187. Dorsotemiinal seta of legs, hollow, pilose

Fig. 188. (Chemo) sensory seta, blunt

Fig. 189. (Chemo) sensory seta, attenuate
Figs. 190-193. Modifications of peglike setae

Fig. 194. Duplex seta, insertions contiguous

Fig. 195. Duplex seta, insertions approximate

Fig. 196. Duplex seta, insertions separate

Fig. 197. puplex seta with microseta modified as a hollow peg

A Revision of the Mite Family Bdellidae

499

Figures 178-197

179

80 181 182 183 184

191 193

i) i i (b

185 186 187 188 189 190 192

194

197

27-8920

Publications of the University of Kansas Science Bulletin

Kansas University Quarterly

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