'JV-

OCCASIONAL PAPERS
OF THE

California Academy of Sciences

No. 124, 51 pages, 120 figures.

TYPE SPECIMENS OF TIPULIDAE DESCRIBED BY R. W. DOANE IN THE
COLLECTION OF THE CALIFORNIA ACADEMY OF SCIENCES

By

George W. Byers

Department of Entomology

University of Kansas

Lawrence, Kansas 66045

Abstract: The entomological career of R. W. Doane
is briefly reviewed. For each included species, the
following are presented: name (original combination),
date and page reference to original description, pre-
sent taxonomic status, numbers and kinds of types,
condition of types, and comments. Most holotypes and
lectotypes are illustrated. Lectotypes are designated
for Diaranomyia adjeata , D. particeps , Pachyrhina
oatifoTnioa I P. snowii , Rhypholophus arcuatus / R.
Gornutus , R. longiaornus , R. paratlelus , Tiputa alia,
T. aspersa , T. atvisumma , T. biuncus , T, aalijovnioa ,
T. colovadensis , T. cylindvata , T. derbyi , T. flavo-
oauda, T. flavomarginata , T. madera, T. marina, T.
newcomevi , T. ooaidentalis , T. pacifioa , T. plani-
oornis , T. pyramis , T. quay Hi , T. vohweri , T. vusti-
Gola, T. silvestra , T. sternata , T. sylvicola , T.
tergata , T. ungulata , T. vestigipennis , T. williamsii ,

T". Contribution no. 1600 from the Department of Entomology,
University of Kansas, Lawrence, Kansas 66045.

Q

^^' viU

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Rennie Wilbur Doane is known primarily as an economic
entomologist, the author of several books in applied ento-
mology, including Insect Pests (1910), Eoonomic Entomology
and Zoology (with V. L. Kellogg, 1915) , Insects and Disease
(1925), and Forest Insects (with E. C. VanDyke , W. F. Cham-
berlin and H. E. Burke, 1936). However, early in his pro-
fessional career he became interested in Tephritidae and
particularly Tipulidae, and for over a decade (1900-1912) he
published extensively on the taxonomy of Tipulidae, describ-
ing in a dozen papers 151 species, mostly from the western
United States. The bibliography that follows lists all of
Doane 's publications on Tipulidae.

Doane was born in Des Moines, Iowa, in 1871. At Leland
Stanford Jr. University, he studied entomology when it was
first offered there, under Prof. John Henry Comstock and
Vernon L. Kellogg. Graduating in 1896, Doane took a position
as Assistant Zoologist and Entomologist at Washington Agri-
cultural College, Pullman. It was here that he began assem-
bling his collection of crane flies. Following his promotion
to Assistant Professor, Doane was assigned in 1901 to the
Fisheries Experiment Station at Keyport, Washington, as its
superintendent. Several of his tipulid specimens bear the
Keyport label, which pertains to an area about nine miles
north of Bremerton, in Kitsap County, Washington, on a chan-
nel of Puget Sound.

In the fall of 1905, Doane returned to California to
accept an appointment as Assistant Entomologist at Stanford
University. Here he was promoted to Instructor and Curator
in 1906, and in 1909 was made Assistant Professor. He
attained the rank of Associate Professor in 1915 and Profes-
sor in 1926. His work on the crane flies did not take him
far afield, and most species described from his own collec-
tions of Tipulidae were from Palo Alto (Stanford) or other
localities in the San Francisco Bay area. His developing
interest in economic entomology, in contrast, took him in
1908 to the Society Islands and in 1913 to Samoa, to bring
coconut pests under control. In California, he worked on the
biology and control of various economic pests, particularly
scale insects, on fleas in relation to plague, and on mos-
quito control. He retired in 1936 and died at Stanford on
1 December 1942. His crane fly collection was subsequently
transferred from Stanford to the California Academy of
Sciences; however, much of the collection assembled while
Doane was associated with Washington Agricultural College
(now Washington State University) remains in Pullman.

Sources of these biographical data are Pierson (1943) ,
Alexander (1967), Essig (1931) and Osborn (1937).

In general, the type specimens of Doane 's species
described in 1908 through 1912 are in the collection of the
California Academy of Sciences (CAS) , while those described
in his earlier papers are in the Washington State University
collection. He seems to have brought with him to Stanford,
however, a few representatives of the species described in

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE

1900 and 1901, and in nearly every instance these may be
identified as syntypes. Lectotypes for the 1900-1901 species
should, where possible, be selected from the holdings at
Washington State University (WSU) .

Usually, Doane based his descriptions of new species on
stated numbers of males and females, or on stated single
specimens. Where there was 9nly one specimen, it is of
course the holotype. Where there were several, Doane usually
in some way labelled only one as "type," and this one was
put into the CAS collection with the status of holotype.
Occasionally two or more syntypes were given "type" labels.
Other original syntypes may or may not have been subsequently
labelled, with a yellow tag, as "PROBABLE/PARATYPE/ (specif ic
name in capital letters)." This segregation of types from
Doane' s collection was done under the direction of Dr. E. S.
Ross in 1946.

Many of the labelled "types" lack genitalia or wings
(generally the right wing) . Presence on certain specimens
of such a label as "Hypo 3" suggests the hypopygium was dis-
sected off and given a code number for subsequent association
with that syntype. In a letter to E. S. Ross, dated 20 May
1946, G. F. Ferris wrote, concerning disposicion of the
Doane collection: "...I know that there is nothing left that
didn't have definite indications as to significance and
origin. He had, for example, a lot of slides of tipulid
wings with no indication of what they came from, so I didn't
keep them. " In selecting lectotypes , I have sometimes by-
passed such an apparently dissected specimen labelled "type"
in favor of a more completely intact specimen from among the
other syntypes. It seems to me most useful to designate as
lectotype of each species the specimen that at present
affords the most information about the species and that is
also consistent with present understanding of the species.

According to the International Code of Zoological Nomen-
clature, Article 73, the presence of these "type" labels
does not alter the status of syntypes, that is, does not
confer on the specimens so labelled the status of holotype
and upon other syntypes the status of paratypes . A holotype
is to be designated in the published description (or must be
the only specimen) .

Acknowledgments

Dr. Paul H. Arnaud, Jr. , Curator and Chairman of the
Department of Entomology, California Academy of Sciences,
originated this re-examination of Doane ' s types as part of a
program to catalogue all types in the Academy's collection.
I am indebted to him for inviting me to San Francisco for
part of June 1974 to study the types, and to him and his wife
Madeline for truly extraordinary hospitality during my stay.
Other members of the Department of Entomology at the Academy
also contributed friendly assistance in many ways. I thank

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Dr. William J. Turner, Director of the M. T. James Entomo-
logical Collection, Washington State University, Pullman,
for his efforts in identifying type specimens among the
Doane materials in his care. Finally, I express appreciation
to Emeritus Professor Charles P. Alexander of Amherst, Massa-
chusetts, for his help to me in this work, and for similar
generous assistance in crane fly matters over the past 25
years .

Format for the Species List

In the treatment of individual species I present infor-
mation in the following sequence:

1. Name of the species in its original combination ,
followed on the same line by

2. Author's name (Doane) and

3. Date of publication of the description (with letter
if necessary to indicate sequence of papers within a single
year) , and

4. Inclusive pages of description of the species.

5. Present status of the species, indicating such
things as change of generic assignment, current subgeneric
assignment, or synonymy.

6. Enumeration of the original type series and state-
ment of the type locality or localities.

7. Portion of the type series now present in the collec-
tion of the California Academy of Sciences (CAS) .

8. Lectotype designation, if appropriate, including
pin-label data for the lectotype (or similar information for
the holotype) . Inscriptions on each pin-label are given
within quotation marks, with diagonal lines separating lines
on the label.

9. Condition of the holotype or lectotype.

10. Taxonomic comments, including such things as recog-
nition characteristics of the species, references to figures,
comparison of figures with others already published, notes
on similar species, synonymy, etc.

Species are arranged in alphabetical order within genera;
the genera are similarly in alphabetical sequence.

Abbreviations

Follov/ing are abbreviations used repeatedly on the
plates: adm - adminiculum, bs - basistyle, gon - gonapo-
physis, id - inner dististyle, p - proctiger (tenth abdominal
segment) , 8s - eighth abdominal sternum, 9s - ninth abdominal
sternum, 9t - ninth abdominal tergum.

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE

Amalopis ampla Doane , 1900: 195.

Present status: Vedicia (Trioyphona) ampla ampla
(Doane) .

Based on Q6 , 19 from Seattle, Washington. CAS collec-
tion: 16 syntype. Apparently only Ic^ , 19 syntypes remain
in the WSU collection.

Amalopis oonctans Doane, 1900: 196.

Present status: Pedieia (Trioyphona) oonstans (Doane).

Described from 6 or more specimens representing both
sexes, from Olympia (restricted type locality), Seattle,
Tokeland and South Bend, Washington. CAS collection: 16
syntype, from Tokeland, Washington. There are 26 and a
broken specimen from Olympia in the WSU collection.

Diaranomyia adjecta Doane, 1908a: 8.

Present status: Limonia (Hespevolimonia) infuseata
(Doane) .

Doane based the species on 26 , 29 from the vicinity of
Stanford University (Santa Clara Co. ) , California. CAS
collection: 16, 29 syntypes.

LECTOTYPE , by present designation, the male syntype (CAS
#5632), labelled "Corte Ma-/dera Cr. Cal./ 5/15-1906"; "R.
W. Doane/Col lector " and "Di cranomyi a/adj ecta/07 Doane" (in
pencil, Doane ' s handwriting). A label pinned beside this
specimen reads "Presumed type from head of Doane series
(E.S.R. 1946)." All the legs and the left antenna are
missing from the lectotype.

Alexander (1967: 85) gave the date of collection of one
of the female syntypes as 12 October 1907, suggesting there
might be a second generation each year; however, the label
actually reads April (IV) . There is another male in the CAS
collection without date that could possibly be the missing
syntype, but it bears a printed label similar to that of an
adjacent specimen dated 1909 and is therefore not considered
part of the type series .

The illustration by Alexander (1967: 231, fig. 253) is
good, so I have not provided a further sketch of the lecto-
type. It should perhaps be noted that the base of the
ventral dististyle is expanded somewhat dorsad, producing a
shallow cleft in which the base of the dorsal dististyle
lies. Also, the downturned, blackened spines at the apex of
the dorsal dististyle are noticeably blunt- tipped.

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Dicranomyia oervina Doane , 1908a: 8-9.

Present status: Limonia ( Diaranomyia) cervina (Doane).

Described from a single male from the vicinity of Stan-
ford University (Santa Clara Co.), California. CAS collec-
tion: the holotype.

HOLOTYPE, male (CAS #5633), labelled "Woodside Cal/ 4/4
1906"; "R. W. Doane/Collector"; "Di cranomy i a/cervi na/07
Doane" and "N.B. = distans O.S./C.P.A." The specimen is
generally intact but lacks the right wing and left hind leg,
and the genital segments have been removed to a microscope
slide.

Alexander (1965: 46) regarded cervina as a synonym of
distans (Osten Sacken) . The two forms are similar in general
appearance, wing characters, and even hypopygial structure.
But in oervina there is a conspicous , thumb-like projection
from near the base of the rostrum of the ventral dististyle
(figs. 1,2), the rostrum of the dististyle is distinctly
rounded at apex, and the rostral spines are broader, more
blade-like than those of distans. Comparison of my illus-
trations of cervina with Alexander's (1966: 317, fig. 35C)
suggests also that cervina has a more curved dorsal disti-
style and lacks a conspicuous ventral process on the basi-
style (but the latter may be concealed by the slide pre-
paration) .

Doane 's published generic name, Dicranomsia , was clearly
only a typographical error.

Dicranomyia particeps Doane, 1908a: 7-8.

Present status: Limonia (Dicranomyia) particeps (Doane).

The species was described on the basis of 96, 6?, all
from Keyport (Kitsap Co.), Washington. CAS collection: 4c5 ,
49, 2 without abdomen, all syntypes.

LECTOTYPE, by present designation, a male (CAS #5631) ,
labelled "7" (on a small tag); "Keyport Wash/July 1905 RWD"
and "particeps." The specimen lacks both hind legs, and its
terminal abdominal segments have been mounted on a micro-
scope slide (see Alexander, 1959: 49).

One male from the type series was sent to C. P. Alex-
ander; and since one of the syntypes without abdomen bears
a label reading "Hypo 3," it is assumed to be a male. Four
syntypes are apparently lost. The lectotype is selected to
conform to Alexander's differentiation of particeps and
ypsilon (Alexander, 1959). Of the four male syntypes present
in the CAS collection, however, three are Limonia ypsilon
Alexander, including the most fully labelled male from the
head of the series as it stood in Doane 's collection.
Doane' s description applies as well to particeps as to
ypsilon, and it seems to me that the present designation of

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE

a lectotype does away with what might have become a taxonomic
problem.

At the time he described L, ypsilon , Alexander (1959)
supposed that his Limonia ( Dioranomyia) uinta (Alexander,
1948: 38-40) was a synonym of L. (D.) partioeps (Doane) .
After comparing details of the lectotype of particeps (figs.
3, 4) with Alexander's (1948) illustration of uinta, however,
I regard the two forms as distinct. Particularly the mesal
lobe of the basistyle and the arrangement of setae on the
rostrum of the ventral dististyle differ between these
species .

Diavanomyia vividioans Doane, 1908a: 7.

Present status: Limonia (Dicranomyia) humidicola (Osten
Sacken) .

Described from H from the vicinity of Stanford Univer-
sity (Santa Clara Co.), California. CAS collection: the
holotype.

HOLOTYPE, male (CAS #5630) , labelled "Corte Ma-/clera Cr
Cal/2 1906"; "R. W. Doane/Col 1 ector " and "Di cranomya/ vi ri di -
cans/07 Doane" (Doane's misspelling). The holotype is a
teneral specimen lacking left antenna, left middle leg and
right front and middle legs. Its genital segments have been
removed to a microscope slide; the left dististyles are
shown in fig. 5.

Erioaera austera Doane, 1900: 192-193.

Present status: Hexatoma (Erioaera) austera (Doane).

Species based on lUi from Olympia, Washington. CAS
collection: 16 labelled only "4 25/94"; "R. W. Doane/Col 1 ec-
tion" and "Probabi e/ Paratype/aus tera " with an identification
label, in Doane's pencilled writing, "Eri ocera/aus tera/
Doane," This is probably a syntype. The WSU collection has
four males similarly lacking locality data yet probably syn-
types .

Paahyrhina breviorcornis Doane, 1908b: 178-179.

Present status: Nephrotoma breviorcornis (Doane).
Described from a single m.ale from Battle Creek (Calhoun
Co.), Michigan. CAS collection: the holotype.

HOLOTYPE, male (CAS #5637), labelled "Battle Creek/
F^ich."; "type" (in pencil); and "Pachy rhi na/brevi orcorni s/
Doane." it lacks the right wing, apical half of the left
antenna, and all legs except the right femur and part of the
tibia. In the box with the type is a leg that was found

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

loose with the specimen in 1970.

Recognition characteristics of this species include its
13-segmented antennae, form of ninth abdominal tergum (fig.
6), and long, curved, horn-like gonapophyses (fig. 7, gon) .

Pachyrhina oalifornica Doane , 1908b: 176.

Present status: Tiputa ( Hesperotipula) oalifornica
(Doane) .

The original description was based on "many males and
females" from Stanford University (Santa Clara Co.), Cali-
fornia. CAS collection: 19, 1 mating pair and possibly
other syntypes , and 16 labelled "type" but not accepted as
such, as discussed below.

The specimen bearing the "type" label is a damaged male
also labelled "San Jose Cal/Apr. 1906" and "Pachyrhina/
cal i forni ca/02 type Doane." Both the type labels have more
than one pin hole so may have been transferred from some
other specimen. Since this specimen has a locality label
not in agreement with the published type locality, I do not
accept it as the type or as a syntype. Alexander (1967: 29)
designated as lectotype a male from Stanford University,
stating it was in the CAS collection. It seems likely he
did not see the specimen labelled as the "type." The only
male specimens in the CAS collection from "Stanford Univer-
sity" were collected in April 1908, the month of publication
of the species' description. A female collected at Stanford
University on 26 April 1904 and a mating pair labelled "Palo
Alto Cal./6 April '95" might be parts of the original type
series .

LECTOTYPE. In view of Alexander's designation of a
lectotype, I have selected a male, intact except for lacking
the right front and hind legs, to be lectotype (CAS #5634).
It is labelled "Stan U Cal/20 Apr 1908" and "R. W. Doane/
Collection," the date suggesting it may not actually be part
of the original type series (unless publication of the April
1908 issue of Entomological News was considerably delayed) .

In the original description, the generic name was inad-
vertently spelled Paahyrina.

Pachyrhina oocidentalis Doane, 1908b: 177-178.

Present status: Nephrotoma wulpiana (Bergroth) .

Described on the basis of "many males and females" from
localities in California, Montana, Washington, Nevada and
Arizona. But Doane specified that the type (by which he
apparently meant to differentiate a holotype, in the current
sense, from a series of paratypes - or at least intended to
restrict the type locality) was from Stanford (Santa Clara
Co.), California. CAS collection: the "type" male and 46,

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE

109 "paratypes." There are also Id, 3? and 4 without abdo-
men, all probable "paratypes," in the WSU collection.

HOLOTYPE, male (CAS #5636), labelled "Stanf o rd/ Uni v .
Cal"; "type" and " Pachy rh i na/occi den ta 1 i s/Doane . " It lacks
the right wing, most of the right antennal flagellum, and
all legs (but one leg is loose in the box with the specimen) .

This commonly collected western species apparently
broadly overlaps in range the similar Nephrotoma fevruginea
(Fabricius) of more eastern North America but is the only
representative of the genus in the Pacific coast states and
British Columbia. The ninth abdominal tergum (fig. 9) and
the dense brush of golden hairs on the margins of the eighth
sternum (fig. 8) will differentiate males of wulpiana from
those of furruginea . Concerning the illustration: right
lateral aspect is shown because the left side of the type is
obscured by the wing; and the attenuated apex of the outer
dististyle is curved downward out of sight.

Pachy vhina snowii Doane, 1908b: 176-177.

Present status: Nephrotoma snowii (Doane).
Description based on 16, 1? from Lander (Fremont Co.),
Wyoming. CAS collection: only the male syntype.

LECTOTYPE, by present designation, the male syntype (CAS
#5635), labelled "Near Lander Wyo./5000 to 8000 ft. July/Roy
Moodie"; "24" (in red ink); and "Pachy rh i na/s nowi i / Doane . "
The lectotype lacks its right wing, both antennae, and all
legs except the right front one and part of the right middle
femur.

Alexander (1965) divided snowii into a typical subspecies
from Wyoming and a subspecies altevnata (Dietz) from Colo-
rado.

General aspects of the terminal abdominal segments of
the lectotype are shown in figures 10 and 11. In dorsal
aspect, the ninth tergum is broadly rounded (nearly semi-
circular) , shallowly indented medially, its posterior margins
bearing many small, black denticles most densely crowded
beside the indentation, or notch.

iV. snowii resembles in many details ¥1 . vittula (Loew,
1864) and may prove to be a synonym, of that species.

Rhypholophus avcuatus Doane, 1908c: 201.

Present status: Ormosia (Ormosia) avcuata (Doane).
This species was based on 29 from Ithaca (Tompkins Co.),
New York. CAS collection: both syntypes ,

LECTOTYPE, by present designation, a female syntype (CAS
#5638), labelled "Ithaca, N. Y./24 Apr. '94" and "Rhypholo-
phus/arcuatus/Doane/Type . " The right wing and all legs

10 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

except the right hind one are missing. Wing length is 7.35
mm, not 8 mm as described. Figure 12 is the ventral aspect
of the left wing of the lectotype.

Rhypholophus oornutus Doane , 1908c: 202.

Present status: Ormosia (Ormosia) cornuta (Doane) .
Description based on 26, 19 from Stanford (Santa Clara
Co.)/ California. CAS collection: all three syntypes.

LECTOTYPE, by present designation, a male (CAS #5642) ,
labelled "Corte Ma-/dera Cr Cal/ 5/15 1906"; "R. W. Doane/
Collector"; "R. W. Doane/Col 1 ecti on" and "Probabl e/Paratype/
cornutus." This male has no legs or antennal flagella
remaining, while the female syntype , which had been labelled
as "type," is more intact. Since specific identification in
Ormosia depends upon characters of the male genitalia, how-
ever, I have designated as lectotype the syntype that has
most taxonomic utility. The second male has had the genital
segments removed to a microscope slide, it appears (i.e., is
labelled "Hypo 24"), and that slide can no longer be found.
The lectotype (figs. 13, 14) is in agreement with figure 483
in Alexander's Crane Flies of California (1965: 263).

Rhypholophus divexus Doane, 1908c: 201.

Present status: Ormosia (Parormosia) fusiformis fusi-
formis (Doane) .

Described from a single female specimen from Keyport
(Kitsap Co.), Washington. CAS collection: the holotype.

HOLOTYPE, female (CAS #5639), labelled "Keyport Wash/
July 1905 RWD" and "Rhyphol ophus/di vexus/Doane/Type . " The
specimen lacks right wing, tip of right antenna, and all
legs except right middle one and left front femur. The
discal cell in the holotype is abnormal, being partially
open into cell 2nd M2 , but the specimen agrees with the
description of fusiformis in other respects. I have illus-
trated the left wing in ventral aspect (fig. 15) to facili-
tate comparison with illustrations of other species.

Rhypholophus longicornus Doane, 1908c: 201-202.

Present status: Ormosia (Ormosia) longicornis (Doane)
Original spelling emended by Alexander (1965: 86).

Described on the basis of 26 , 29 from Keyport (Kitsap
Co.), Washington. CAS collection: all four syntypes.

LECTOTYPE, by present designation, a male (CAS #5640),
labelled "Keyport Wash/July 1905 RWD"; "slide #1" and

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 11

"Rhyphol ophus/1 ongi cornus/Doane/Type . " The genital segments
(fig. 18) have been removed to a microscope slide, which is
present with the type. Apical two-thirds of the right
antenna are missing. The antennae of the male (fig. 16) are
long and conspicuously verticillate , if extended parallel to
the costa reaching nearly to the tip of Sc (cf. fig. 17).

Rhypholophus parallelus Doane , 1908c: 202.

Present status: Ormosia (Ormosia) parallela (Doane).
Described from 2? from Ithaca (Tompkins Co.), New York.
CAS collection: both syntypes.

LECTOTYPE, by present designation, a female syntype (CAS
#5641), labelled "Ithaca, N. Y." and " Rhyphol ophus/ paral -
1 el us/Doane/Type. " The lectotype lacks its right wing, left
antennal flagellum and half of right flagellum, and all legs
except the right middle and hind ones. By use of the key to
Ormosia provided by Alexander (1966: 462-464), pavallela can
be grouped with three or four other described species. It
seems to me similar to 0. rubella (Osten Sacken) . Species
in Ormosia have been differentiated largely on the basis of
male genitalia, however, and it is probably best to leave
parallela as an unrecognizable form (Alexander, 1965) until
the genus is examined in detail. My accompanying figures 19
and 20 of the lectotype do not seem to me to be in any way
diagnostic but might give some information to a better
taxonomist .

Tipula acuta Doane, 1901: 116-117.

Present status: Tipula (Triplioitipula) acuta Doane.

Described on the basis of 46, 49, apparently all from
Palo Alto (Santa Clara Co.), California. CAS collection:
39 syntypes, all from Palo Alto. The WSU collection con-
tains 3c5 , 19 and one without abdomen, probably all syntypes

Tipula acutipleura Doane, 1912: 42.

Present status: Tipula (Lunatipula) acutipleura Doane.
Described from a single male from San Diego (San Diego
Co.), California. CAS collection: the holotype.

HOLOTYPE, male (CAS #5648), labelled "San Diego/Cal.";
''Type" and "Ti pul a/acu tapl eura/Doane/ ' 1 1 . " (The misspelling
is Doane ' s . ) It is intact except that the left front leg,
right middle leg, and parts of two tarsi are missing.

In the holotype, the lobes of the ninth abdominal tergum.
(fig. 23) are less sharp than figured by Alexander (1967:
191, fig. 119), and the adminiculum (fig. 22, adm) seems

12 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

less complicated when not slide-mounted. Characteristic of
this uncommon species are the large inner dististyles and
acutely prolonged basistyles (fig. 21) and yellowish,
pubescent cushions on the mesal, posterior margins of the
ninth sternum (fig. 22).

Tipula aequalis Doane , 1901: 108.

Present status: Tipula (Triplicitipula) aequalis Doane.

The species was based on 19c^ , 49 from various places in
Washington and California, with Pullman, Washington, the
restricted type locality. CAS collection: Ic^ syntype from
Keyport, Washington. There remain only A6 , 19 in the WSU
collection.

Tipula albimaoula Doane, 1912: 51.

Present status: Tipula (Bellavdina) albimaoula Doane.
Described from one male from an unspecified locality in
Arizona. CAS collection: the holotype.

HOLOTYPE, male (CAS #5662), labelled "Arizona/C. U. Lot
35"; below this a small rectangle of bluish paper without
inscription; and "Ti pul a/ a 1 bi macul a/Doane/ ' 1 1 . " The speci-
men has had the apical one-third of its abdomen cut off, and
the right wing and all legs but the right hind one are
missing.

The Rs is only slightly longer than the m-cu cross-vein.
One striking characteristic of this species is the long
verticils of the antennae, these being more than twice the
length of their respective f lagellomeres near the base of
the flagellum (fig. 24).

Tipula alia Doane, 1911: 161-162.

Present status: Tipula (Beringotipula) fallax Loew.

The type series included 9i , 29 from Keyport (Kitsap
Co.), Washington, and 19 from Olympia (Thurston Co.), Wash-
ington. CAS collection: 2<i , 29 syntypes , all from Keyport.

LECTOTYPE, by present designation, a male (CAS #5643),
labelled "Keyport Wash/July 1905 RWD" and "R. W. Doane/Col-
lection." This specimen is intact except for the loss of
the left middle leg. The male labelled "Hypo in vial /Key 1" ;
"wing photo 2" and "Type" was not selected because the
separated hypopygium cannot be located at present. The left
wing, which is attached to a card beneath the specimen, was
illustrated by Doane in ventral aspect. The third male has
its abdomen intact but is otherwise considerably damaged.

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 13

At the time he described T. alia, Doane differentiated
it from T. fallax Loew. Specimens in his collection identi-
fied as T. fallax, however, are not that species. Alexander
examined the lectotype of T. fallax in 1964 and subsequently
illustrated that species in detail (1967: 179, fig. 76),
showing three distinct branches , or arms , on the pendulous
appendage of the inner dististyle. Doane (1911: 166) had
supposed there were only two branches. Specimens lacking
the prominent, hook-like middle portion of the appendage
seem in other respects similar to T. neweomevi Doane (com-
pare my figs. 76, 79 and Alexander's 1967 fig. 81). I am
indebted to Dr. Alexander for a series of slides of T. new-
comeri showing variation in this structure. Some additional
details of the terminal abdominal segments of the lectotype
of T. fallax Loew are included here (figs. 25-27) for com-
parison with T. newcomeri and related species of subgenus
Bevingotipula. It should be noted that the outer dististyle
in figure 25 is curved mesad apically so that its rounded
margin is concealed.

The specific name was misspelled olia in the description
but was spelled correctly elsewhere in the paper and was
corrected in the errata for volume 18 of Psyche.

Tipula alta Doane, 1912: 44.

Present status: Tipula (Fterelaohisus ) alta Doane.

Described on the basis of a single male individual from
near Lander (Fremont Co.), Wyoming. CAS collection: the
holotype.

HOLOTYPE, male (CAS #5651), labelled "Near Lander Wyo . /
5000 to 8000 ft. July/Roy Moodie"; "61" (in red ink); "Type"
and "Ti pul a/al ta/Doane/ ' 1 1 . " The specimen is intact with
respect to body and wings but lacks all legs except the left
front one.

This is a small Tipula. The open discal cell noted by
Doane is probably an anomaly. The hypopygium is character-
ized by large, complicated inner dististyles (figs. 28, 29),
a small pubescent lobe at the posterior apex of each side of
the ninth sternum, and a caudally facing, small, dark,
polished concavity beneath the margin of the ninth tergum
(concealed in dorsal aspect, fig. 30).

Tipula aspersa Doane, 1912: 51-52.

Present status: Tipula. ( Sinotipula) aspersa Doane.

This species was based on 16 , 19 from Pacific Grove
(Monterey Co.), California. CAS collection: the male syn-
type.

14 CALIFORlNlIA ACADEMY OF SCIENCES [Occ. Papers

LECTOTYPE , by present designation, the male syntype (CAS
#5663), labelled "Pac 6r Cal/30 Oct 1906" and "T i pul a/asper-
sus/Doane/ ' 1 1 . " The lectotype lacks the apex of the left
antenna and all legs from the left side (but one leg is
loose in the tray with the specimen) .

Alexander (1967: 215, fig. 197) showed the ninth abdomi-
nal tergum with rather broad apical lobes tipped with numer-
ous black denticles - the appearance of the tergum when
cleared and slide-mounted. In the dried lectotype, the
lobes are slender, with divergent tips, the denticles on the
lower surfaces and not visible from above. Figure 31 shows
these differences as well as the characteristic pattern of
distribution of the short hairs of the tergum. The poste-
rior aspect of the hypopygium (fig. 32) may add somewhat to
understanding the complex dististyles (cf. Alexander's fig.
197) .

There is in the Doane collection (CAS) a female that was
pinned next to the lectotype until E. S. Ross in 194 6 segre-
gated the types. It may be the female syntype but is
labelled "Corte Madera Cr., Cal . , 10-22-1907," which differs
from the published type locality.

Tipula atrisumma Doane, 1912: 42-43.

Present status: Tipula (Lunatipula) atrisumma Doane.
Species based on llc5 , 59 from San Diego (San Diego Co.),
California. CAS collection: 5c^ , 59 syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5649), labelled "San Diego/Cal."; "Type" and "Ti pul a/atra-
summa/Doane/ 1 91 1 . " The lectotype is intact except for
lacking its left hind leg.

Alexander (1967: 40) emended the specific name to atra-
summa, as it was spelled on the pin label. In my opinion,
however, this is an unjustified emendation under the terms
of the International Code of Zoological Nomenclature. Fur-
ther, it seems to me that Doane meant the name to say "black
tip," for which atrisumma would be a correct term. Figures
33-35 present some aspects of the lectotype not shown by
Alexander (1967: 193, fig. 125).

Tipula biarmata Doane, 1912: 55.

Present status: Tipula (Lunatipula) albo fascia Doane.
Described on the basis of Ic^ from Keyport (Kitsap Co.),
Washington. CAS collection: the holotype.

HOLOTYPE, male (CAS #5669), labelled "Keyport Wash/July
1905 RWD"; "Type" and "Ti pul a/bi armata/Doane/1 1 . " The apex
of the right wing and both front and middle legs are missing
from the holotype.

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 15

The illustration by Alexander (1967: 191, fig. 121) por-
trays the ninth abdominal tergum in a composite of dorsal
and ventral aspects, and the processes of the inner disti-
style somewhat out of their natural positional relationships,
probably as a result of use of a slide-mounted specimen.
Since this species {T. albo fascia) is very similar to T.
bifaloata in many details, I have provided sketches of taxo-
nomically useful aspects of the hypopygium (figs. 36-39) for
comparison with Alexander's illustrations of T. bifaloata.

Tipula bifaloata Doane , 1912: 55.

Present status: Tipula (Lunatipula) bifaloata Doane.
Described from a single male taken at San Diego (San
Diego Co.), California. CAS collection: the holotype.

HOLOTYPE, male (CAS #5668), labelled "San Di ego/Cal . " ;
"Type" and "Ti pul a/bi f a1 cata/Doane/ ' 1 1 . " The holotype lacks
its abdomen beyond the third segment and all its legs except
the left hind one. It has little taxonomic usefulness, but
in those characteristics that can be compared it seems to
agree with Alexander's (1967: 34, 191, fig. 122) identifi-
cation of the species.

Tipula bisetosa Doane, 1901: 111-112.

Present status: Tipula (Lunatipula) bisetosa bisetosa
Doane .

Description based on 19'^, 14? syntypes from Pullman,
Washington, and 2d syntypes from Collins, Idaho. CAS collec-
tion: lc5 , 19 syntypes from Pullman. In the WSU collection
are 3c5 , 29 and 2 syntypes without abdomen.

Tipula bituberoulata Doane, 1901: 101.

Present status: Tipula (Tvi oho tipula) bituberoulata
Doane.

Species described from 26 , 19 from an unstated locality
in California. The WSU collection has Id syntype. CAS col-
lection: 2d, labelled "San Diego, Cal . " and "Probable/Para-
type/bi tubercul ata . " Since the WSU syntype bears the type
number assigned by Doane (#153) , it is clear that not both
males in the CAS collection are syntypes, unless Doane had
more specimens than he indicated in the description.

16 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Tipula biunous Doane , 1912: 58.

Present status: Tipula (Lunatipula) biunous Doane.

The species was described on the basis of H , 1? from an
unspecified locality in southern California. CAS collection:
both syntypes.

LECTOTYPE, by present designation, the male syntype (CAS
#5673), labelled "S. Cala."; "C. U. Lot 35"; below this a
blank slip of bluish paper; "Type" and "Ti pul a/bi uncus/
Doane/ '11." The lectotype lacks the front and hind legs on
both sides and the apical one-third of the right antennal
f lagellum.

Alexander (1965: 35) recently catalogued the species
under its original name but later (Alexander, 1967: 41)
apparently inadvertently spelled the name biunoa.

To my knowledge, this species has never been illustrated.
Accordingly, some taxonomically important details of the
lectotype are included here (figs. 40-42). The ninth abdom-
inal tergum (fig. 41) is characterized by two somewhat diver-
gent, blunt, darkened lobes and a U-shaped median notch
flanked by thin, nearly transparent marginal blades. Two
darkly sclerotized tips, apparently of the adminiculum, pro-
ject conspicuously backward (fig. 42). These structures,
from which Doane derived the specific name, lie in a verti-
cally oriented fork that seems to be another apex of the
adminiculum (fig. 40).

Tipula oalifovnica Doane, 1912: 49.

Present status: Tipula (Triplicitipula) doaneiana Alex-
ander. The specific name oalifovnioa is preoccupied in
Tipula by oalifovnioa Doane, 1908 (described as a Faohyrhina) .

Described on the basis of 26 specimens from Palo Alto
(Santa Clara Co.), California. CAS collection: both syn-
types ,

LECTOTYPE, by present designation, a male syntype (CAS
#5660), labelled "Palo Alto, Cal./2 Apr. '92" (or possibly

'97, the year pencilled over a printed '93) and "Tipula/
cal i f orni ca/Doane/1 91 1 . " The specimen lacks all right legs,
left middle and hind legs, and knob of the right haltere.
The paralectotype lacks its terminal abdominal segments; it
is more distinctly dated 14 Apr. '92 (over '93).

Large spines on abdominal sternum 8 shown by Alexander

(1967: 223, fig. 229) are not visible in the types, probably
being concealed by dense hairs. Also concealed in my drawing

(fig. 44) is the apex of the adminiculum ( "phallosome" ) ,
which in the lectotype has the membranous tenth segment
resting upon it. A hypopygial character not mentioned by
earlier authors is a broad, thick, transverse flange on the
lov/er ninth sternum (figs. 43, 44). Against the posterior

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 17

face of this flange, on each side, hangs a brush of fine,
coiled golden hairs.

Ti-pula carinata Doane , 1901: 103-104.

Present status: Tipula (Platytipula) carinata Doane.

Species based on 456 , 129 syntypes from Pullman (Whitman
Co.), Washington. CAS collection: 16, 19 syntypes. There
are 126, 29 remaining in the WSU collection.

Tipula oognata Doane, 1901: 123-124.

Present status: Tipula (Yamatotipula) cognata Doane.

Described from 69 syntypes from Seattle and Olympia,
Washington, and Moscow Mountain (near Moscow) , Idaho. It
appears to have been Doane 's intent to restrict the type
locality to Olympia (Thurston Co.), Washington. CAS collec-
tion: one of 5d that Doane only tentatively assigned to this
species, thus not technically a syntype , although it was be-
fore Doane when he described T. cognata.

Tipula coloradensis Doane, 1911: 164-165.

Present status: Tipula (Beringotipula) coloradensis
Doane.

The original description was based on 16 , 39 from Taber-
nash (Grand Co.), Colorado. CAS collection: all four syn-
types, as well as 16 with same locality and date labels but
technically not part of the type series.

LECTOTYPE, by present designation, the male syntype (CAS
#5646), labelled "Tabernash ,/ Col . Aug./E. S. Tucker"; "L. S.
Jr. U./Lot 492/Sub. 6"; "Hypo in/vial"; "Type" and "Tipula/
col oradensi s/Doane/ 1 91 1 . " This specimen lacks the apical
one-third of the abdomen, the right haltere , and all legs
except the left hind femur. It appears that Doane removed
the hypopygium, but this can no longer be found. The second
male in the CAS collection is a topotype, bearing the first
two labels listed for the lectotype. On the basis of this
more intact male (figs. 46-50), I judge that Tipula fundata
Alexander (1945: 407), listed by Alexander (1965: 32) as a
synonym of T. coloradensis , is a valid species. There are
clear differences in the shape of the apex of the ninth
abdominal tergum, the gonapophyses , and especially the inner
dististyles.

18 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Tipula oylindrata Doane , 1912: 46.

Present status: Tipula (Serratipula) eylindrata Doane.

The species was described on the basis of 3<^ from San
Diego (San Diego Co.), California. CAS collection: 3c5 syn-
types .

LECTOTYPE, by present designation, a male syntype (CAS
#5655), labelled "San Diego/Cal."; "Type" and "Ti pul a/cyl i n-
drata/Doane/ 1 91 1 . " The lectotype lacks its left front and
middle legs and right hind leg, has some psocid or dermestid
damage to the thorax, and had the right wing almost torn off
near the base (the last now repaired) . One of the paralecto-
types is in slightly better condition than the lectotype but
is not labelled "Type."

A characteristic of this species not mentioned in exist-
ing descriptions is the arrangement of hairs on the terminal
abdominal segments. On the ninth tergum and sternum and
most of the eighth sternum there is a covering of fine hairs
bent cephalad at their bases and closely appressed to the
surface. These hairs are of such fineness, density and
nearly parallel arrangement as to give to the surface a look
of brushed satin. Scattered among them are larger hairs,
more erect yet still inclined cephalad (contrast my figs. 51,
53 and fig. 190 of Alexander, 1967). In the types, the
posterior margin of the ninth tergum is glabrous, and the
lateral lobes are not distinctly truncate (cf. Alexander,
1967: 213, fig. 190). The ninth sternum is deeply cleft,
nearly to the posterior margin of the eighth sternum, which
is unmodified (fig. 52) .

Tipula devbyi Doane, 1912: 47-48.

Present status: Tipula (Hesperotipula) derhyi Doane.

Original description based on "many males, 6 females,"
all from Stanford University (then Palo Alto, Santa Clara
Co.), California. CAS collection: llOci, 399 having label
data suggesting they might belong to the type series, as dis-
cussed below. From these, 1Q16 and 6? are selected, on the
basis of condition, to comprise the syntypic series.

LECTOTYPE, by present designation, a male syntype (CAS
#5658), labelled "L. S. Jr. U./Lot 517/Sub 34" and "R. W.
Doane/Col 1 ect i on . " The male earlier labelled as "Type" is
not selected because of its generally poor condition; the
lectotype is the best preserved male in the series, taking
into consideration easy visibility of the genital structures.

There are three major groups of specimens potentially
comprising the type series. One, consisting of 54c5 , 129, is
labelled "Stan U Cal/20 Apr 1908" and "R. W. Doane/Col 1 ec-
tion." Two females in this group have been labelled "Prob-
able/ Pa ra type/ derby i " by someone on the CAS curatorial staff;

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 19

these, therefore, are accepted as two of the six intended
female syntypes. The second group comprises 5 3ci , 229,
labelled "L. S. Jr. U./Lot 517/Sub 38" and "R. W. Doane/Co1-
lection." One male from this group bears the "Type" label
and Doane ' s identification label. From this group I have
selected the two females in best condition to be part of the
syntypic series. The third group, 26, 59, is labelled "L. S.
Jr. U./Lot 517/Sub 34" and "R. W. Doane/Col 1 ecti on . " One
female in this group bears Doane 's pencilled "Type" label
(as on the male mentioned) ; it and the best remaining female
in the group are accepted as the last two female syntypes,
completing the six. There seems to be no reason to reject
any df the males in these groups from syntypic status, except
perhaps that some are no longer taxonomically useful speci-
mens. Accordingly, 106 of them are regarded as paralecto-
types .

Alexander (1967: 185, fig. 102) illustrated the compli-
cated inner dististyle, eighth sternum and ninth tergum. In
my sketch of the lateral aspect of the lectotype (fig. 54) ,
I have omitted the inner dististyle in order to show more
clearly the profile of the ninth tergum. The dorsal aspect
of the ninth tergum is shown in figure 55.

Tipula dovsolineata Doane, 1901: 98.

Present status: Tipula (Triohoti-pula) dovsolineata
Doane.

Based on 5c5 , 109 syntypes from Pullman, Washington, and
19 syntype from Keyport, Washington. Because of his peculiar
method of indicating restriction of type locality, Doane 's
intent has, I think, been mistaken by Alexander (1967: 59),
who took the Keyport female to have the status of holotype.
By comparison of locality data for T. dovsolineata with those
for T. illustris on the same page, one sees that the word
"Type" follows the intended typical locality designation.
Moreover, Doane regularly selected a male as "Type" (or so
labelled the specimens) unless he had only females before
him. CAS collection? 16 (without abdomen), 19 fairly
intact, and 19 without abdomen, all from Pullman,
Washington. In the WSU collection are 26, 19 syntypes, all
from Pullman.

Tipula flaviaoma Doane, 1912: 57.

Present status: Tipula (Lunatipula) incisa incisa Doane.
Described from a single male from Montana, without fur-
ther locality data. CAS collection: the holotype.

HOLOTYPE, male (CAS #5672), labelled "Montana/C. U. Lot
35"; a blank slip of bluish paper; "Type" and "Tipula/f lavi-
coma/Doane/ ' 11 . " The specimen lacks its right wing and apex
of the left wing (beyond stigma and cell 1st M2) and all legs

20 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

except the left hind one.

Tipula incisa is a widespread western species. The typi-
cal subspecies was described from Washington by Doane in
1901. It occurs also in British Columbia and Montana. Other
subspecies are recognized from New Mexico, Arizona, Kansas
and Oklahoma, and as far east as Ohio (Alexander, 1965: 36).

Some details of the hypopygium of the holotype of T.
flavicoma are shown in figures 56-58.

Tipula flavocauda Doane, 1912: 60.

Present status: Tipula ( Lunatipula) flavocauda Doane.

Description based on 3cJ , 3? , said in the original de-
scription to be from San Diego, California, but labelled
"Arizona." CAS collection: 3cJ , 2? syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5677), labelled "Arizona/C. U. Lot 35."; "Ill"; a blank
slip of bluish paper; "Type" and "Ti pul a/fl avocauda/ Doane/
1911." The specimen lacks the right front and middle legs,
left middle leg, and knob of right haltere.

Tipula (L. ) flavocauda has been recorded three times from
San Bernardino Co., California (Alexander, 1967: 42), so it
may actually occur at or near the erroneously published type
locality. Alexander (1965: 36; 1967) listed T. buenoi as a
synonym of T. flavocauda. Although this seems probable, I
note differences especially in the ninth abdominal tergum
and sternum between the lectotype of T. flavocauda (figs.
59-62) and the specimen of T, buenoi illustrated by Alexander
(1946: 500, fig. 4) .

Tipula flavomarginata Doane, 1912: 46-47.

Present status: Tipula (Lunatipula) flavomarginata
Doane.

Described from 86 from San Diego (San Diego Co.), Cali-
fornia. CAS collection: 26 syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5656), labelled "San Diego/Cal."; "R. W. Doane/Coll ecti on "
and "Probabl e/Paratype/f 1 a vomargi nata . " This male lacks the
left middle leg, right front and middle legs, and most of the
left and half of the right antennal flagella. Another male
is labelled "Type" but is not made lectotype because it is
even more damaged and because the genital segments are not
in taxonomically useful condition.

Since the illustration of the ninth abdominal tergum
given by Alexander (1967: 195, fig. 133) is a composite of
dorsal and ventral aspects, I have provided a dorsal view of
this sclerite in the lectotype (fig. 64). The lateral aspect
(fig. 63) shows some taxonomically useful details of the

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 21

ninth sternum, as well as the length of the ventral projec-
tions from the ninth tergum.

Tipula fulvinodus Doane , 1912: 45.

Present status: Tiputa (Luyiatipula) fulvinodus Doane.
Species described from a single male from Grand Coulee
(Grant Co.), Washington. CAS collection: the holotype.

HOLOTYPE, male (CAS #5653), labelled "Grand Coulee, Wash/
Foster Coulee. /24. June '02"; "Type" and "Ti pul a/ful vi nodus/
Doane/ '11." Body and wings of the holotype are intact, but
the right front, left middle and both hind legs are missing.

This species will not be identified as a Lunatipula by
existing keys because it lacks the tuft of small setae on
the squama of the wing, as noted by Alexander (1948: 283).
In other respects, though, it appears to belong to that sub-
genus .

Some details of the hypopygium of the holotype are shown
in figures 65 and 66.

Tipula fumosa Doane, 1901: 99-100.

Present status: Tipula ( Yamato tipula) dejecta Walker.

This species was based on 76 , 19 syntypes from Columbus
(Franklin Co.), Ohio. CAS collection: 26 syntypes, one of
them without abdomen. There is 16 syntype in the WSU collec-
tion.

Tipula illustris Doane, 1901: 97-98.

Present status: Tipula (Angarotipula) illustris Doane.

Described on the basis of 16 syntypes from Idaho, Wash-
ington, and Michigan. Doane apparently intended to restrict
the type locality to St. Anthony (Fremont Co.), Idaho. CAS
collection: 16 syntype, without abdomen, from Olympia, Wash-
ington. Three males remain in the WSU collection.

Tipula impudioa Doane, 1901: 104.

Present status: Tipula (Lunatipula) impudioa Doane.

Based on 56 , 19 syntypes from various localities in Wash-
ington, the restricted type locality being Wawawai , Whitman
Co. CAS collection: IcJ syntype from Almota, Washington,
and 16, 19 syntypes from Pullman, Washington. Two additional
males are in the WSU collection.

22 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Tipula inoisa Doane , 1901: 118.

Present status: Tipula ( Lunatiputa) inoisa inoisa Doane,
Based on 4d , 19 syntypes from Wawawai and 29 syntypes
from Pullman (all Whitman Co.), Washington. CAS collection:
Irf syntype from Wawawai, without its abdomen. There are 26 ,
29 in the WSU collection. Doane also described this species
as T. flaviooma; see under that name, above.

Tipula inouvva Doane, 1912: 43.

Present status: Tipula (Pterelaohisus ) inouvva Doane.

The species was described from one male specimen from an
unspecified locality in Nebraska. CAS collection: the holo-
type.

HOLOTYPE, male (CAS #5650), labelled "Nebraska"; "V 24
01"; "Hypo in/vial"; "Type" and "Ti pul a/ i ncurva/ Doane/ ' 1 1 . "
It lacks the abdomen, all legs, left antennal flagellum and
right haltere.

Alexander (1965: 33) supposed T, inouvva might be a
synonym of T. angulata Loew , but this seems unlikely to me.
Although the holotype is in rather dilapidated condition, it
can be placed with confidence in the tvivittata group of sub-
genus Ptevelaohisus . Of particular significance is the ab-
sence of the nasus , a characteristic not noted by Doane. I
think T. inouvva may be a western Nebraska species.

The right wing of the holotype is shown in figure 67.

Tipula lamellata Doane, 1901: 105-106.

Present status: Tipula (Lunatipula) lamellata Doane.

Description based on 56 , 19 syntypes from Pullman (Whit-
man Co.), Washington. CAS collection: 36 syntypes, two of
them without abdomen. The remaining 26 , 19 syntypes are in
the WSU collection.

Tipula madeva Doane, 1911: 162-163.

Present status: Tipula (Bevingotipula) madeva Doane.

Species described on the basis of A6 , 29 from Stanford
University and vicinity (Santa Clara Co.), California. CAS
collection: 26, 29 syntypes.

LECTOTYPE , by present designation, a male syntyoe (CAS
#5644), labelled "Stan U Ca1/28 May 1908"; "R. W. Doane/
Collection" and "Ti pul a/madera/Doane/ ' 1 1 . " The lectotype
lacks all its legs except the right hind one and has the
right antennal flagellum broken near mid-length but adhered
to the left flagellum. Otherwise it is in good condition.

Wo. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 23

Another male, from Corte Madera Creek, near the Stanford
campus, has been labelled "Type" but is not selected as
lectotype because it lacks the terminal abdominal segments
and thus is of limited taxonomic utility.

Alexander (1967: 179, fig. 78) gives a composite dorsal-
ventral illustration of the ninth abdominal tergum. The
median-apical process of this tergum is best seen in postero-
dorsal aspect (fig. 69). Figures 68 and 70 present further
details of the hypopygium of the lectotype.

Tipula marina Doane , 1912: 44-45.

Present status: Tipula ( Sevvatipula) marina Doane.
Described from 3c5 syntypes from Palo Alto (Santa Clara
Co.), California. CAS collection: 2d syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5652), labelled "Salt Marshes/Palo Alto Cal/ 3/22 1906";
"R. W. Doane/Col 1 ector" ; "Type" and "Ti pul a/mari na/Doane/
'11." This specimen is intact except for the lack of its
right hind leg.

The median tuft of hairs on the posterior margin of the
eighth abdominal sternum, illustrated by Alexander (1967:
213, fig. 192), is not evident on the lectotype. Also, the
posterolateral corners of the ninth tergum in the lectotype
(fig. 72) are more rounded than shown by Alexander, and the
inner dististyle (fig. 71) is of a somewhat different appear-
ance, in part due to the fact that the abundant hairs of its
dorsal and mesal surfaces are virtually recumbent, rather
than erect as shown by him.

Tipula meridiana Doane, 1912: 58.

Present status: Tipula (Yamatotipula) mevidiana mevidi-
ana Doane.

Described on the basis of a single male from Arizona,
without more detailed locality. CAS collection: the holo-
type.

HOLOTYPE, male (CAS #5674), labelled "Arizona/C. U. Lot
35."; a small bluish tag without inscription; "Type" and
"Ti pul a/meri di ana/Doane/ ' 1 1 , " This specimen lacks all legs
except part of the right front femur, but one leg is glued
to the pin-label. Both antennae are broken off at the base.

Alexander (1965: 28) lists the range of this form as
Arizona, California and Utah, the California record being
from Monterey Co. (Alexander, 1967: 69). The range of sub-
species oontinentalis is given as Alaska to California and
Colorado, with records from many parts of California. Thus,
the ranges of these two nominal subspecies overlap broadly.
As I am at present unaware of all the evidence, I am

24 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

following Alexander's placement of the two forms. Since the
evidence from ranges clashes with my concept of subspecies,
however, it seems to me that subspecific rank might have
been assigned on the basis of degree of difference and that
T. meridiana Doane and T. aontinentalis Alexander are more
likely similar but different species.

In the holotype, the ninth abdominal tergum (figs. 74,
75) is less widely cleft and its apical lobes less covered
with blackened denticles than shown by Alexander (1967: 229,
fig. 245) . There is a conspicuous projection between abdom-
inal sterna 8 and 9 (fig. 73) .

Tipula newaomeri Doane, 1911: 163-164.

Present status: Tipula (Beringotipula) newcomeri Doane.
Original desciption based on Ad , 4V from Deer Park
(Placer Co.), California. CAS collection: AS, 4? syntypes,
as well as four additional specimens with label data that
indicate they were collected together with the types.

LECTOTYPE , by present designation, a male syntype (CAS
#5645), labelled "Deer Park/Placer Co./Cal./El. 6500/Jun. 25
'08"; "Type" and "Ti pul a/newconieri /Doane . " The lectotype is
generally intact, but the right front and hind legs and left
middle leg are missing, as well as the apical two-thirds of
the right antennal flagellum.

This seems to differ from the species illustrated as T.
newaomeri by Alexander (1967: 24, 178, fig. 81). The pendu-
lous lobe of the inner dististyle more nearly resembles that
shown by Alexander (1967: 178, fig. 76) for T. fallax, which
is otherwise quite different in structure. The gonapophyses
of the lectotype and male paralectotype (fig. 77) are as
shown by Alexander for T. newaomeri , but there are conspicu-
ous differences in the shapes of the outer dististyle and
apical armature of the ninth abdominal tergum (cf. my figs.
76, 78, 79).

Although there are 66 , 59 and one broken specimen from
Deer Park (June and July, 1908 and 1909) in the Doane collec-
tion, and I can see no reason to exclude any of them from
the type series, I have selected the 4(5 and 49 in best con-
dition to reconstitute the described series of eight indi-
viduals.

Tipula nigroaorporis Doane, 1912: 45.

Present status: Tipula (Vestiplex) nigroaorporis Doane,
Described on the basis of one male from Estes Park
(Larimer Co.), Colorado. CAS collection: the holotype.

HOLOTYPE, male (CAS #5654), labelled "Estes Park/August
'92/F. H. Snow"; "27" (in red ink); "Type"; "Tipula n sp./

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 25

goes to serta Lw. but/color etc not right" and "Ti pul a/ni gro-
corpori s/Doane/1 91 1 . " The holotype lacks all its legs, but
part of one is glued to the upper pin-label. The specimen
is otherwise generally intact.

The coloration of this species is striking: head,
thorax and first abdominal segment are blackish brown to
black, with areas of silver-gray pollinosity. The rest of
the abdomen is nearly orange (Doane called it brownish yel-
low) , with brownish lateral margins on the terga. The ninth
tergum (fig. 81) bears an irregular transverse ridge near
mid-length and has the unevenly rounded lateral lobes trans-
versely grooved posteriorly; its postero-median area merges
with the dorsum of the membranous tenth segment (proctiger) .
Most of the antennal f lagellomeres are deeply indented ven-
trally, with a thick ventral lobe just beyond the base (fig.
80) .

Tipula oocidentalis Doane, 1912: 59-60.

Present status: Tipula (Triplioitipula) pvaeoisa Loew.
New synonymy.

Description based on 6cJ specimens from San Diego (San
Diego Co.), California. CAS collection: 6cJ syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5676), labelled "San Diego/Cal."; "Type" and "Ti pul a/occi -
dental i s/Doane/1 91 1 . " The lectotype has all legs except
the right middle one missing but is otherwise in good con-
dition.

Alexander (1967: 64) considered T. oocidentalis "very
close to pvaecisa Loew" and thought the two "may prove to be
synonymous." It is, in fact, curious that Doane described
this species since in his collection are several specimens
identified by him as T. pvaecisa Loew that are clearly the
same as his T. occidentalis . I have compared my notes and
sketches (figs. 82-84) of the lectotype of T. occidentalis
with the somewhat larger male syntype of T. pvaecisa Loew,
and in my opinion both specimens represent the same species.

In the CAS collection are 16 specimens with label data
(place) same as on the lectotype. Five of these have been
labelled "Probabl e/Paratype/occi dental i s . " These are
accepted as the five besides the lectotype that comprise
the described type series, though there seems no particular
reason to exclude any of the seven.

Tipula pacifica Doane, 1912: 48-49.

Present status: Tipula (Sinotipula) pacifica Doane.

The original description was based on 3c^ , 29 from Deer
Park, Placer Co. , California, and 19 from Keyport (Kitsap
Co.), Washington. CAS collection: H, 29 syntypes from

26 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Placer Co., California, and 19 from Keyport, Washington.
Doane recorded the California specimens as "Types," restrict-
ing the type locality, but the Washington female has syntypic
status .

LECTOTYPE, by present designation, the remaining male
syntype (CAS #5659), labelled "Deerpark/Pl acer Co. /El. 6500
Cal./Aug. 1, 1908"; "Type" and "Ti pul a/paci f i ca/Doane/ 1 91 1 . "
The specimen lacks its right front and middle legs and has
the right rear leg broken off and reattached with glue. A
female labelled the same as the lectotype for locality and
date is also labelled "type 9" in Doane ' s writing. The
second female from California is labelled only "Placer Co./
VII-29-09/E1 . 8500 ft." and not Deer Park.

Although existing illustrations of this species (Alex-
ander, 1967, figs. 203-205) are generally accurate, certain
additional details from the lectotype may be seen in my
figures 85 and 86.

Tipula planicornis Doane, 1912: 52-53 (as planicornia) .

Present status: T-iputa (Tviplicitiputa) ptaniaornis
Doane.

The type series consisted of 20<^ , 89 from San Diego (San
Diego Co.), California. CAS collection: 12<^ , 29 syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5664), labelled "San Diego/Cal."; "R. W. Doane/Col 1 ecti on "
and "T. pi ani corni s/D" (in Doane's pencilled writing). The
lectotype is intact except for the lack of its right front
leg.

The original spelling, planicovnia , is improperly formed,
and the emendation by Alexander (1967) seems justified under
Article 33 of the International Code of Zoological Nomen-
clature. It seems also from the manuscript identification
label that planicovnis was the name Doane intended. Only
two specimens in the type series bear name labels in Doane's
writing, and on both labels the termination of the specific
name is "-is. "

The similarity in hypopygial structure between T. plani-
covnis (Figs. 87-89) and T. vestigipennis (Figs. 116-117) is
striking, but so is the difference in wing development. See
comments under T. vestigipennis .

Tiputa pyvamis Doane, 1912: 53.

Present status: Tipula (Eremotipula) pelluaida Doane.
Alexander (1965: 38) stated this synonymy but subsequently
(1967: 26) apparently inadvertently omitted it. It seems to
me correct, on the basis of published figures of T. pellucida ,
although I have not seen the type of T. pellucida (new name

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 27

by Doane , 1912, for T. olava Doane , 1901, preoccupied by T.
olava Kirby, 1884) .

Tipula pyramis was described on the basis of Id syntypes
from Pyramid Lake (Washoe Co.), Nevada. CAS collection: 5ci
syntypes.

LECTOTYPE , by present designation, a male syntype (CAS
#5665), labelled "Pyramid Lake/Nev. June 1911"; "Coll. E. J./
Newcomer"; "R. W. Doane/Col 1 ecti on " and "Probabl e/Paratype/
pyramis." This specimen (figs. 90-92) is intact except for
the loss of its left hind leg and minor repair to the right
front leg. The right haltere is glued to the upper pin-
label. Originally another male syntype was labelled "Type"
but is not selected as lectotype because its hypopygium has
been removed and lost, and there is damage to the legs and
one wing.

Tipula quaylii Doane, 1909: 18-19.

Present status: Tipula (Tviplioitipula) quaylii Doane.

The type series was stated to comprise 15c? , 29 from Yuba
City (Sutter Co.), California. CAS collection: 116, 19
specimens judged to be most of the original series, for
reasons given below.

LECTOTYPE, by present designation, a male (CAS #12074) ,
labelled "L. S. Jr. U./Lot 517 Sub 36"; "R. W. Doane/Collec-
tion" and "Probable/Paratype/quaylii . " This individual is
intact except for missing its left middle leg. The left
hind leg is glued to the first label. The entire series of
116, 19 bear the Leland Stanford Jr. University, Lot 517,
Sublet 36 label. Another male is also labelled "Probable/
Paratype/quaylii" and one other male is labelled "T. quayli/
Yiiba Co." Note the variant spelling of the specific name
and the locality, just east of Sutter Co. I have made
several inquiries to determine if there is any key or
catalogue by which the Stanford University lot numbers might
be identified, but to date I have had no success in this.
The types of T. quaylii should be in Doane ' s Stanford
collection, and I am assuming that the series discussed here
are the types, sublet 36 referring to Quayle's collection
from Yuba City (or Yuba County) .

In this small, brownish gray Tipula the wings are shorter
than the halteres (figs. 95-96). The ninth tergum and some
other parts of the abdomen (figs. 93-94) are covered with
fine, pale, appressed hairs that give them a satin-like
lustre.

2 8 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Tipula retusa Doane , 1901: 109.

Present status: Tipula (Lunatipula) retusa Doane.

Described from 10c5 , 49 syntypes from various localities
in Washington. CAS collection: Id syntype from Wawawai,
24 April 1897, and 19 syntype from Seattle, without date.
Remaining in the WSU collection are 26 , 39 syntypes and one
broken specimen that is probably a syntype.

Tipula rohweri Doane, 1911: 165.

Present status: Tipula (Beringotipula) rohweri Doane.

Described from 36, 19 syntypes from Florissant (Teller
Co.), Colorado, and 26, 39 from Big Horn County, Wyoming.
CAS collection: 26, 19 syntypes from Florissant and lc5 , 19
syntypes from Big Horn County.

LECTOTYPE, by present designation, a male syntype (CAS
#5647), labelled "Florissant/Colo. July/26 (Ckll.)"; "in
marsh"; "Type" and "Ti pul a/rohweri /Doane/1 91 1 . " The apical
half of the abdomen is missing from the lectotype , as well
as the apical half of each antennal flagellum, the right
hind leg and left middle and hind legs. The right wing has
been removed to a card that accompanies the lectotype, and
there is one leg loose in the tray beside the type. The
second male from Florissant is damaged even more and is not
of much taxonomic use. Only the male from Big Horn Co. ,
Wyoming, has the genital segments intact. Doane indicated
that Florissant, Colorado, was the type locality, but since
he seems to have had a good eye for structural details, I
think we can accept the Wyoming male (figs. 97-99) as repre-
sentative of the species. Although the Colorado and Wyoming
localities are separated by nearly 400 miles, the respective
specimens seem to be conspecific with regard to characters
that can be compared.

Tipula (B. ) monoana Alexander seems to differ only
slightly from T. rohweri in structure of the hypopygium.

Tipula rupioola Doane, 1912: 50.

Present status: Tipula (Bellardina) rupioola Doane.
Described from a single male taken in Oak Creek Canyon
(Coconino Co.), Arizona. CAS collection: the holotype.

HOLOTYPE, male (CAS #5661), labelled "Oak Creek Canon/
Ariz. 6000 ft. Aug/F. H. Snow"; "57" (in red ink) and
"Ti pul a/rupi col a/Doane/ ' 1 1 . " The apex of the abdomen has
been cut off at the fifth segment, and the specimen lacks
the right wing, left hind leg and right front and middle
legs.

Since the radial sector is slightly longer than the m-cu

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE 29

cross-vein, this individual would run to subgenus Sinotipula
by existing keys. Doane described in considerable detail
the hypopygium that is now missing.

Tipula rusticola Doane, 1912: 47.

Present status: Tipula (Setitipula) rustioola Doane.
This species was based on 26 specimens from Keyport
(Kitsap Co.), Washington. CAS collection: both syntypes.

LECTOTYPE , by present designation, a male syntype (CAS
#5657), labelled "Keyport Wash/July 1905 RWD"; "Type" and
"Ti pul a/ rus ti col a/Doane/ 1 91 1 . " The lectotype lacks right
middle leg and both hind legs and has a 2-3 mm section of
the costa of the right wing, just before the stigma, missing,
But it is generally in good condition. The other male lacks
the apical portion of its abdomen.

The ninth abdominal tergum (fig. 100) has notched outer
lobes and a shallow median depression without hairs. The
complexity of the inner dististyles is shown better by Alex-
ander (1967: 213, fig. 195) than by my figure 101, yet the
latter shows the conspicuous lateral projection from the
posterior portion of the dististyle. The gonapophyses are
fringed with large, flattened setae (fig. 101), and there is
a whitish median zone extending from the bases of the gonapo-
physes to the unmodified posterior edge of the eighth ster-
num, which is also whitish.

Tipula silvestra Doane, 1909: 18.

Present status: Tipula (Triplioitipula) silvestra Doane.

Described from 29 specimens taken at Pacific Grove
(Monterey Co.) and 19 from Stanford University (Santa Clara
Co.), California. CAS collection: both syntypes from
Pacific Grove.

LECTOTYPE, by present designation, a female syntype (CAS
#12075), labelled " I 1 1 - 29- 08/ Pac . Grove, /Cal." and "R. W.
Doane/Col 1 ecti on . " The lectotype lacks all legs on the left
side and the right hind leg but is otherwise intact.

Although not bearing any earlier type labels, these two
females from Doane ' s collection are regarded as two of the
three syntypes. Their label data are correct. No other
specimens are present in the Doane collection.

The structure of the ovipositor seems taxonomically help-
ful, the eighth sternum especially being long, its hypovalves
increasing slightly in width toward the tip and being nearly
truncate at apex (fig. 102). The tenth tergum, separated
from the cerci by an extensive membranous region, is trans-
versely grooved near its posterior end and is 1.45 times as
long as the cerci. Including the hypovalves, the eighth
sternum is 3.0 times as long as the cerci (all measurements

30 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

are greatest length, lateral aspect).

Doane described the male of this species as Tipula ungu-
lata (which see, below).

Tipula simplex Doane, 1901: 103.

Present status: Tipula (Triplicitipula) simplex Doane.

Based on 26 syntypes taken at Palo Alto (Santa Clara
Co. ) , California, and stated by Doane to be in the WSU col-
lection. There are no type specimens in the CAS collection
(cf. Alexander, 1967: 66), and only one of the syntypes
remains in the holdings of WSU. Numerous specimens in the
Doane materials in the CAS are either undated or are dated
after the species was described.

Tipula spatha Doane, 1912: 59.

Present status: Tipula (Lunatipula) spatha Doane.
Species described from one male from an unspecified
locality in Arizona. CAS collection: the holotype.

HOLOTYPE, male (CAS #5675), labelled "Arizona/C. U. Lot
35."; a small, blank, bluish tag; "Type" and "Ti pul a/spatha/
Doane/ '11." The holotype is intact except for the lack of
all legs but the left middle one.

Since, to my knowledge, this species has never before
been illustrated, I offer drawings of details of the hypo-
pygium of the holotype (figs. 103-105).

Tipula spectabilis Doane, 1901: 120-121.

Present status: Tipula (Lunatipula) macrolahis maoro-
labis Loew.

Described on the basis of 10c5 syntypes from Collins
(Bingham Co.), Idaho. CAS collection: 26 syntypes, labelled
"Collins, Idaho/27 July, 1898"; "R. W. Doane/Coll ecti on " and
"Probabl e/Paratype/spectabi 1 i s . " Only one male syntype re-
mains in the WSU collection.

Tipula sternata Doane, 1912: 56.

Present status: Tipula (Lunatipula) sternata Doane.

Species based on 26 from Stanford University (Stanford,
Santa Clara Co.), California. CAS collection: one male
syntype.

LECTOTYPE, by present designation, the remaining male
syntype (CAS #5670), labelled "Stan U Cal/Apr 190-"; "Type"
and "Tipul a/sternata/Doane/ ' 1 1 . " The specimen lacks its

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 31

front legs (except part of the left femur) , parts of the
middle tarsi, and its abdomen, which is broken off at the
thorax. It thus has slight taxonomic value; however, the
species is readily recognizable. The accompanying illus-
trations (figs. 106-107), made from a specimen collected in
Monterey Co., California, 30 May 1950, by J. Linsdale, are
offered to complement those by Alexander (1967: 203, fig.
160) .

Tipula streptooera Doane , 1901: 113.

Present status: Tipula (Hesperotiputa) streptooera
Doane .

This species was based on 46 , 19 syntypes from Collins
(Bingham Co.), Idaho, and Olympia and Tokeland, Washington.
CAS collection: 16 (without abdomen) from Collins, Idaho.
There is also 1? from Collins, Idaho, 27 July 1898. But
since the WSU collection has 16 and 19 from Collins, as well
as the males from Olympia and Tokeland, the female in the
CAS collection cannot be a syntype , despite the label indi-
cating it was collected the same place and date as the types

Tipula sulphurea Doane, 1901: 99.

Present status: Tipula (Yamato tipula) sulphurea sul-
phurea Doane.

Based on 1S6 , 69 syntypes from Battle Creek (Calhoun
Co.), Michigan. CAS collection: 1<^ , 19 syntypes labelled
"Battle Creek/Mich." and 16, probably a syntype, but without
locality label. In the WSU collection, there remain only
2(5 and 1 broken syntypes.

Tipula sylvioola Doane, 1912: 53-54.

Present status: Tipula (Triplioitipula) sylvioola Doane.

Description based on 5c5 syntypes from Keyport (Kitsap
Co.), Washington, and IcJ from an uspecified locality in
southern California. Doane indicated that the type locality
was Keyport. CAS collection: 56 syntypes, including the
one from California.

LECTOTYPE, by present designation, a male syntype (CAS
#5666), labelled "Keyport Wash/July 1905 RWD"; "Type" and
"Ti pul a/syl VI cola/Doane/1 91 1 . " The lectotype is generally
intact, lacking only its right front leg.

The syntype from southern California closely resembles
T. williamsii Doane from the San Francisco Bay area, but T.
williamsii is more darkly marked and has a somewhat more
angular margin to the outer dististyle and paler hairs on
the inner, triangular lobe of the eighth abdominal sternum.

32 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

The darker markings of T. williamsii include especially
wider, darker brownish areas at the sides of the abdominal
terga. Alexander (1967: 66) regarded T. sylvicola as
"allied to aequalis but may be distinct." Specimens identi-
fied as T. aequalis Doane in Doane's collection, however, I
judge to be very nearly the same as, if not conspecific with,
T. praeoisa Loew, but lacking a distinct dorsal brown stripe
on the abdominal terga they will key (Alexander, 1967: 61)
to T. lygvopis Alexander. These specimens are at the same
time quite unlike Alexander's (1967: 223, fig. 227) illus-
tration of T. aequalis with respect to the form of the ninth
abdominal tergum.

Since this species was not illustrated either by Doane
(1912) or Alexander (1967) , I offer three sketches of hypo-
pygial details based on the lectotype (figs. 108-110).

Tipula tergata Doane, 1912: 56-57.

Present status: Tipula (Lunatipula) tergata Doane.

This species was described from two males from Pyramid
Lake (Washoe Co.), Nevada, and one male from an unspecified
locality in southern California. Doane restricted the type
locality to Pyramid Lake. CAS collection: the 26 syntypes
from Nevada.

LECTOTYPE, by present designation, a male syntype (CAS
#5671), labelled "Pyramid Lake/Nev. June 1911"; "R. W. Doane/
Collection"; "Probabl e/Paratype/ tergata " and "T. tergta
Doane/type removed from/here by E. S. R. 1946." This syn-
type lacks all legs except part of the right hind femur but
is otherwise intact. The other syntype, originally labelled
"Type," has had the abdomen removed at the third segment and
lacks three of its legs; therefore, it is not selected as
lectotype.

The hypopygium of the lectotype differs in several de-
tails, notably in the shape of the ninth abdominal tergum,
from the only published illustrations of this species (Alex-
ander, 1967: 203, fig. 161). Accordingly, I have illustrated
it in lateral aspect (fig. Ill) and the ninth tergum in dor-
sal aspect (fig. 112).

Tipula ungulata Doane,- 1912: 54.

Present status: Tipula (Triplicitipula) silvestva Doane.

1 . ungulata was described on the basis of 16<J , 49 syn-
types from San Diego (San Diego Co.), California. CAS col-
lection: 9c?, 19 syntypes.

LECTOTYPE, by present designation, a male syntype (CAS
#5667), labelled "San Diego/Cal."; "Type" and "Ti pul a/ungu-
1 ata/Doane/1 91 1 . " The specimen lacks both front legs but is

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE 33

otherwise intact.

The illustration by Alexander (1967: 227, fig. 239, as
T. silvestra) shows the inner dististyle correctly, the
ninth tergum apparently in ventral aspect (compare with my
fig. 115) , but omits the "strongly chitinized, claw-like
appendages" of the ninth sternum that give the species its
name. These appendages while curved are not sharply tipped,
thus perhaps not strictly claw-like; however, they are con-
spicuous in both the lateral and posterior aspects of the
male hypopygium (figs. 113-114).

Tipula unicincta Doane , 1901: 115-116.

Present status: Tipula (Lunatipula) unicincta Doane.

Description based on 16 , 79 syntypes from Collins (Bing-
ham Co.), Idaho (restricted type locality); Moscow Mountain,
Idaho; Pullman, Washington; and Keyport, Washington. CAS
collection: 16 syntype from Keyport, Washington. There are
36 syntypes and 2 without abdomen in the WSU collection.

Tipula vestigipennis Doane, 1908d: 47.

Present status: Tipula (Tviplioitipula) vestigipennis
Doane.

Described from A6 , 39 from San Francisco (San Francisco
Co.), California, and 19 from Stanford (Santa Clara Co.),
California. CAS collection: 26, 39 syntypes.

LECTOTYPE, by present designation, a male (CAS #12076),
labelled "San Francisco/ 5/7 ' 07/Wi 1 1 i ams " (written in pen-
cil); "L. S. Jr. U./Lot 517/Sub 30"; "R. W. Doane/Col 1 ecti on " ;
"Probable/Paratype/vestigipennis" and "Tipul a/vest ipennis/
Doane/D." (Note Doane ' s spelling error.) The lectotype
lacks its right wing and right hind leg but is generally in-
tact.

This species is structurally very similar to Tipula
planioornis Doane, except that the wings are extremely re-
duced. (Compare my figs. 116-117 and 87-89.) It is known
only from the vicinity of San Francisco Bay, while T. plani-
cornis has so far been found only farther south in Califor-
nia. The possibility that these two nominal species are
geographic races of a single species needs to be investigated.

Tipula williamsii Doane, 1909: 19.

Present status: Tipula (Tviplicitipula) williamsii
Doane .

In the original description, Doane stated that the type
series consisted of 5<J , 69 specimens from San Francisco (San

34 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Francisco Co.)/ California. CAS collection: 56, 69 speci-
mens judged to be syntypes , as discussed below.

LECTOTYPE, by present designation, a male (CAS #12077) ,
labelled "L. S. Jr. U./Lot 517/Sub 37"; "R. W. Doane/
Collection"; "Probable/Paratype/williamsii" and "Tipula/
williamsii/Doane/D. " This specimen is intact except for
lacking its right front and both hind legs.

The CAS collection includes under T. williamsii a series
of 33cJ , 499 with identical label data, i.e., sublet 37 of
lot 517 in the Stanford University collection. Since the
types of williamsii should be in the Doane (Stanford) col-
lection and are not known to be elsewhere, I am assuming
that they are among this lot. Accordingly, I have selected
the S6 , 69 in best condition to make up the type series , and
have labelled them appropriately. It seems likely that F. X,
Williams gave Doane additional specimens after learning that
they represented an undescribed species.

See comments under Tipula sylvioola ! above.

Alexander (1967: 227, fig. 242) illustrated the inner
dististyle in apparently somewhat mesal aspect (cf. my fig.
118) , and his drawing of the eighth sternum shows the sub-
triangular median lobe, which is concealed in my figure 120.
He also emended the specific name to williamsi .

References

ALEXANDER, C. P.

1946. Records and descriptions of North American crane-
flies (Diptera) . Part VI. Tipuloidea of Ari-
zona, New Mexico and Trans-Pecos Texas, I.
American Midland Naturalist, vol. 35, pp. 484-
531.

1948. Records and descriptions of North American crane-

flies (Diptera). Part VII. The Tipuloidea of
Utah, I. American Midland Naturalist, vol. 39,
pp. 1-82.

1949. Records and descriptions of North American crane-

flies (Diptera). Part VIII. The Tipuloidea of
Washington, I. American Midland Naturalist,
vol. 42, pp. 257-333.

1959. Undescribed species of crane-flies from the

western United States and Canada (Dipt. : Tipuli-
dae) . Part XIX. Entomological News, vol. 70,
pp. 47-54.

1965. Family Tipulidae. Pp. 16-90 in Stone, A. et al.,
A catalog of the Diptera of America north of
Mexico. Agricultural Research Service, United
States Department of Agriculture, Handbook No.
276.

No. 124]

BYERS:

TYPE SPECIMENS OF TIPULIDAE

35

1966.

1967.

DOANE, R. W,
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1901.

1907.

1908a.

1908b.

1908c.

1908d.
1908e.

1908f .

1909.

1911.

1912.

ESSIG, E. O,
1931.

OSBORN, H.
1937.

PIERSON, C.
1943.

Family Tipulidae. Pp. 196-486 in Crampton, G.
C. , C. H. Cur ran and C. P. Alexander, Guide
to the insects of Connecticut, Part VI, The
Diptera or true flies of Connecticut. First
fascicle. Connecticut State Geological and
Natural History Survey, Bulletin 64, originally
published 1942, reprinted 1966.

The crane flies of California. Bulletin of the
California Insect Survey, vol. 8, pp. 1-269.

New North American Tipulidae. Journal of the
New York Entomological Society, vol. 8,
pp. 182-198, plates VII, VIII.

Descriptions of new Tipulidae. Journal of the
New York Entomological Society, vol. 9, pp. 97-
127.

Notes on the subapterous female of Tiipula simplex
Doane. Entomological News, vol. 18, pp. 15-16.

Notes on the tipulid genus Dicvanomyia. Entomo-
logical News, vol. 19, pp. 5-9.

New North American Paohyr-hina , with a table for
determining the species. Entomological News,
vol. 19, pp. 173-179.

New species of the tipulid genus Rhypholophus ,
with a table for determining the North American
species. Entomological News, vol. 19, pp. 200-
202.

A new species of Tipula with vestigial wings.
Psyche, vol. 15, pp. 47-49.

Variations in the wing venation in some Tipuli-
dae. Entomological News, vol. 19, pp, 405-407,
plate XVII.

A remarkable outbreak of Tipula larvae. Entomo-
logical News, vol. 19, pp. 437-438.

More Tipula with vestigial wings. Psyche, vol.
16, pp. 17-19.

Tipula fallax and others. Psyche, vol. 18, pp.
160-166.

New western Tipula. Annals of the Entomological
Society of America, vol. 5, pp. 41-61.

A history of entomology. Macmillan Co. , New
York. 1029 pp.

Fragments of entomological history. Columbus,
Ohio; published by the author.

J.

Rennie Wilbur Doane, 1871-1942. Journal of Eco-
nomic Entomology, vol. 36, p. 131.

36

CALIFORI^IA ACADEMY OF SCIENCES [Occ. Papers

<S.w.6.

FIGURES 1 5. FIGURES 1, 2. Dioranomyia aevvina Doane,
male holotype. 1, right dististyles, dorsal aspect. 2,
same, ventral aspect. FIGURES 3, 4. Dioranomyia particeps
Doane, male lectotype. 3, right basistyle, dististyles and
portion of ninth tergum, dorsal asoect. 4, inner lobe of
basistyle, ventral aspect. FIGURE 5. Dioranomyia viridioans
Doane, male holotype, left dististyles, dorsal aspect, most
hairs omitted.

No. 124]

BYERS

TYPE SPECIMENS OF TIPULIDAE

37

G w. 6.

FIGURES 6-11. FIGURES 6, 7. Paahyrhina hreviorcornis
Doane , male holotype. 6, posteromedian portion of ninth
tergum, dorsal aspect. 7, terminal abdominal segments, left
lateral aspect (gon - gonapophysis ) . FIGURES 8, 9. Pachy-
rhina oacidentalis Doane, male holotype. 8, terminal abdom-
inal segments, right lateral aspect. 9, posteromedian por-
tion of ninth tergum, dorsal aspect. FIGURES 10, 11.
Paahyrhina snowii Doane, male lectotype. 10, terminal
abdominal segments, left lateral aspect. 11, hypopygiiom,
posterior aspect. Upper scale: figures 6, 9.

38

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

13

14

16

0

at-

Smrn

1 mm

_i

0

17

G.W. £).

FIGURES 12-20
female lectotype,
Ehypholophus corn
dorsal aspect. 1
rior) aspect. FI
female holotype,
Ehypholophus long
sentative flagell
basistyle and dis
Ehypholophus para
wing. 20, termin
Scale a: figures
Scale c: figures

FIGURE 12. Ehypholophus arcuatus Doane ,
left wing, ventral aspect. FIGURES 13, 14
utus , male lectotype. 13, ninth tergum,
4, left ventral dististyle, ventral (poste-
GURE 15. Ehypholophus divexus Doane,
left wing, ventral aspect. FIGURES 16-18.
icornis Doane, male lectotype. 16, repre-
ar segments. 17, right wing. 18, right
tistyles, dorsal aspect. FIGURES 19, 20.
llelus Doane, female lectotype. 19, left
al abdominal segments, right lateral aspect,

12, 15, 17, 19. Scale b: figures 16, 20.

13, 14, 18.

No. 124]

BYERS

TYPE SPECIMENS OF TIPULIDAE

39

22

FIGURES 21-27. FIGURES 21-23. Tipula acutipleura Doane ,
male holotype. 21, terminal abdominal segments, left lateral
aspect. 22, hypopygium, in part, posterior aspect. 23,
ninth tergum, dorsal aspect. FIGURE 24. Tipula albimacula
Doane, male holotype, basal portion of right antenna, lateral
aspect. FIGURES 25-27. Tipula fallax Loew, male holotype.
25, terminal abdominal segments, left lateral aspect. 26,
lower appendage of inner dististyle, posterolateral aspect.
27, ninth tergum, right lateral aspect. adm - adminiculum,
bs - basistyle, 8s - eighth sternum, 9s - ninth sternum,
9t - ninth tergum.

40

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

C.Vi.B.

FIGURES 28-35. FIGURES 28-30- Tipula alta Doane , male
holotype. 28, terminal abdominal segments, left lateral
aspect. 29, lower parts of hypopygixim, posterior aspect.
30, ninth tergum, dorsal aspect. FIGURES 31, 32. Tipula
aspersa Doane, male lectotype. 31, ninth tergum, dorsal
aspect. 32, hypopygium, posterior aspect. FIGURES 33-35.
Tipula atrisumma Doane, male lectotype. 33, terminal abdom-
inal segments, left lateral aspect. 34, ninth tergum, dor-
sal aspect. 35, eighth sternum, ventral aspect.

No. 124]

BYERS: TYPE SPECIMENS OF TIPULIDAE

41

FIGURES 36-42

le hnloT ia ^^GU^S 36-39. Tipula biarmata Doane ,
leholotype 36, terminal abdominal segments, left lateral
pect 37, ninth tergum, dorsal aspect. 38, posterior

ma
asp

base of inner dististyle, posteromesal aspect. 39, left
naif of apex of eighth sternum, posterior aspect. FIGURES

42

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

43

45

48

44

47

53

FIGURES 43-53. FIGURES 43-45. Tipula oalifovnica
Doane, male lectotype . 43, terminal abdominal segments,
left lateral aspect. 44, hypopygium, posterior aspect.
45, ninth tergum, dorsal aspect. FIGURES 46-50. Tipula
oolovadensis Doane, male lectotype. 46, left dististyles,
left lateral (slightly dorsolateral) aspect. 47, median
process of ninth tergum, posterior aspect, inverted. 48,
ninth tergum, dorsal aspect. 49, inner dististyle and
gonapophyses , posterior aspect. 50, gonapophysis , lateral
aspect. FIGURES 51-53. Tipula cylindvata Doane, male
lectotype. 51, apex of left basistyle and left dististyles,
left anterolateral aspect. 52, ninth sternum, posterior
aspect. 53, ninth tergum, dorsal aspect.

No. 124] BYERS: TYPE SPECIMENS OF TIPULIDAE

43

FIGURES 54-58. FIGURES 54-55. Tiyula derbyi Doane ,
male lectotype. 54, terminal abdominal segments, left
lateral aspect. 55, ninth tergum and adjacent sclerites ,
dorsal aspect. FIGURES 56-58. Tipula flaviooma Doane , male
holotype. 56, terminal abdominal segments, left lateral
aspect. 57, reconstruction of left inner dististyle, lateral
aspect. 58, ninth tergum, dorsal aspect. bs - basistyle,
8t, 8s - eighth tergum and sternum, 9t, 9s - ninth tergum
and sternum.

44

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 59-65. FIGURES 59-62. Tipula flavooauda Doane ,
male lectotype. 59, terminal abdominal segments, left
lateral aspect. 60, ninth tergum, dorsal ^^P^^^ .61 rxght
later-al projection of ninth tergum, to show ventral tooth,
pos?;rio? aspect, enlarged from scale of other f^g^^^s^ H'
lower portions of hypopygium, posterior aspect. FIGURES bJ,
64. Tipula flavomavginata Doane , male lectotype. bJ, ter
minal abdominal segments, left lateral aspect. 64, nmtn
tergum, dorsal asplct. FIGURES 65, 66 T^pu^a /"^^^^^^^^J,,
Doane, male holotype. 65, terminal abdominal segments, left
lateral aspect. 66, ninth tergum, dorsal aspect.

No. 124]

BYERS

TYPE SPECIMENS OF TIPULIDAE

45

C.ViJ.Q

FIGURES ^l-l'o. FIGURE 67. Tipula ino
holotype, right wing. FIGURES 68-70. Tip
male lectotype. 68, left outer dististyle
69, median projection of ninth tergum, pos
inverted, reconstructed in part from paral
left inner dististyle and gonapophyses , po
FIGURES 71, 72. Tipula marina Doane , male
left dististyles, lateral aspect. 72, nin
aspect. FIGURES 73-75. Tipula meridiana
type. 73, terminal abdominal segments, ri
74, ninth tergum, dorsal aspect. 75, apic
ninth tergum, dorsal aspect. Scale a: fi
figures 68, 70-74. Scale c: figures 69,

urva Doane, male
ula madera Doane,
, lateral aspect,
terior aspect,
ectotype. 70,
sterior aspect.

lectotype. 71,
th tergum, dorsal
Doane, male holo-
ght lateral aspect,
al projections of
gure 67, Scale b:
75,

46

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 76-84. FIGURES 16-19. Tipula newaomeri Doane ,
male lectotype. 76, terminal abdominal segments, left
lateral aspect. 77, gonapophysis , lateral aspect. 78,
ninth tergum, posterodorsal aspect- 79, lower appendage of
right inner dististyle, right posterolateral aspect.
FIGURES 80, 81. Tipula nigrooorpovis Doane , male holotype.
80, base of left antenna, left lateral aspect. 81, ninth
tergum, dorsal aspect. FIGURES 82-84. Tipula ocaidentalis
Doane, male lectotype. 82, terminal abdominal segments,
left lateral aspect. 83, ninth tergum, dorsal aspect. 84,
lower portions of hypopygium, posterior aspect.

No. 124]

BYERS

TYPE SPECIMENS OF TIPULIDAE

47

FIGURES 85-92. FIGURES 85, 86. Tipula pa
male lectotype. 85, terminal abdominal segmen
lateral aspect. 86, ninth tergum and apices o
styles, dorsal aspect. FIGURES 87-89. Tipula
Doane, male lectotype. 87, ninth tergum, dors
88, lower portions of hypopygium, posterior as
terminal abdominal segments, left lateral aspe
90-92. Tipula pyramis Doane, male lectotype.
abdominal segments, left lateral aspect. 91,
right basistyle and right dististyles , dorsal
median prolongation of eighth sternum, ventral
adminiculum, bs - basistyle, id - inner distis
eighth tergum and sternum, 9t, 9s - ninth terg

cifica Doane,

ts, left

f outer disti-

planicornis
al aspect,
pect. 89,
ct. FIGURES

90, terminal
ninth tergum,
aspect. 92,

aspect. adm -
tyle, 8t, 8s -
urn and sternum.

48

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

95

96

99

G.w-S,

101

FIGURES 93-101. FIGURES 93-96. Tipula quaylii Doane ,
male lectotype. 93, terminal abdominal segments, left
lateral aspect. 94, ninth tergum, dorsal aspect. 95, left
wing, lateral aspect (cf, fig. 96). 96, left haltere, left
lateral aspect. FIGURES 97-99. Tipula rohwevi Doane, male
lectotype. 97, gonapophysis (shaded) and portion of right
inner dististyle, right lateral aspect. 98, lower appendage
of right inner dististyle, right posterolateral aspect. 99,
median projection of ninth tergum, posterodorsal aspect.
FIGURES 100, 101. Tipula rustiaola Doane, male lectotype.
100, ninth tergum, dorsal aspect. 101, left half of hypo-
pygium, posterior aspect. 9s - ninth sternum. Upper scale:
figures 93-98, 100-101. Lower scale: figure 99.

No. 124]

BYERS

TYPE SPECIMENS OF TIPULIDAE

49

G w.ja.

FIGURES 102-105. FIGURE 102. Tiyula silvestra Doane ,
female lectotype, ovipositor, left lateral aspect. FIGURES
103-105, Tiputa spatha Doane , male holotype. 103, terminal
abdominal segments, left lateral aspect. 104, hypopygium,
posterior aspect. 105, ninth tergima, dorsal aspect. bs -
basistyle, id - inner dististyle, 8t, 8s - eighth tergum and
sternum, 9t, 9s - ninth tergum and sternum.

50

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 106-112. FIGURES 106, 107. Tipula stevnata
Doane, male from Monterey Co., California. 106, terminal
abdominal segments, left lateral aspect. 107, ninth tergum,
dorsal aspect. FIGURES 108-110. Tipula sylvioola Doane ,
male lectotype. 10 8, terminal abdominal segments, right
lateral aspect. 109, ninth tergum, dorsal aspect. 110,
eighth sternum, posterior aspect. FIGURES 111, 112. Tipula
tevgata Doane, male lectotype. Ill, terminal abdominal seg-
ments, left lateral aspect. 112, ninth tergum, dorsal aspect,
bs - basistyle, 8t, 8s - eighth tergum and sternum, 9t, 9s -
ninth tergum and sternum. Scale a: figure 106. Scale b:
figures 107-112.

No. 12 4] BYERS: TYPE SPECIMENS OF TIPULIDAE

51

G. W. B.

FIGURES 113-120. FIGURES 113-1
Doane, male lectotype. 113, terrain
left lateral aspect. 114, hypopygi
115, ninth tergum, dorsal aspect,
vestigi-pennis Doane, raale lectotype
segments, left lateral aspect. 117
aspect. FIGURES 118-120. Tipula w
lectotype. 118, terminal abdominal
aspect. 119, ninth tergum, dorsal
median portions of eighth sternum,
adminiculum, p - proctiger (tenth s
sternum, 9t, 9s - ninth tergum and

15. Tipula ungulata
al abdominal segments,
um, posterior aspect.
FIGURES 116, 117. Tipula

116, terminal abdominal
, ninth tergum, dorsal
illiamsii Doane, male

segments, left lateral
aspect. 120, left and
posterior aspect. adm -
egment) , 8s - eighth
sternum.

COMMITTEE ON PUBLICATION

George E. Lindsay, Chairman
Jean B. Durham, Editor

MBI, WHOI I.IHKARY

UH nCE Y