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HARVARD

Number 1

UNIVERS^Y990

Revision of the Mexican Pyrgomorphidae (Orthoptera: Acridoidea)

II. A reappraisal of the genus Ichthiacris I. Bolivar, 1905, with descriptions of

three new species from Baja California, Mexico.

D. Keith McE. Kevan

Lyman Entomological Museum and Research Laboratory, and Department of Entomology, Macdonald College Campus of
McGill University. 21111 Lakeshore Road. Ste-Anne-de-Bellevue. Quebec H9X ICO. Canada

ABSTRACT.- The orthopteroid genus Ichthiacris Bolfvar (Acridoidea; Pyrgomorphidae) is endemic to NW Mexico. All but one of the species
occur only in Baja California. The genus comprises two subgenera. Ichthiacris. sensu stricto. and Atyphacris Kevan. Singh, and Akbar, 1964.
The latter, formerly regarded as a separate genus, is herein redefined. The genus Ichthiacris now includes eight species: /. (/.) rehni Bolivar
(southern Baja California). /. (/.) par/a n. sp. (known only from Isla Cedros off W Baja California), /. (/.) spinifera n. sp. (southern Baja
California Sur). /. (A.) elongata Kevan et al. (Sonora and Sinaloa). /. (A.) costulata Bolivar (southern Baja California). /. (A.) californica Bolivar
(southern Baja California Sur). /. (A.) celata n. sp. (central Cape region of Baja California Sur), and /. (A.) aptera Hcbard (southernmost Baja
California Sur). The name of the last species reverts from Atyphacris californica (Bruner) to the present name because of recurrent secondary
homonymy. Tabular keys are provided. Biogeographic and phylogenetic relationships are discussed.

RESUMEN.- El genero Ichthiacris Bolivar (Orthoptera: Acridoidea: Pyrgomorphidae) es endemico del noroeste de Mexico. Todas las especies
a excepcion de una de se encuentran solo en Baja California. El genero esta dividido en dos subgeneros, Ichthiacris. sensu stricto. y Atyphacris
Kevan et al., 1964. Este ultimo considerado inicialmente como un genero separado, es redefinido aqui. Los ocho miembros del genero
Ichthiacris ahora considerados son: /. (/.) rehni Bolivar (sur de Baja California), /. (Dparva, n. sp. (conocida solo de lsla Cedros. en la costa del
Baja California oeste). /. (/.) spinifera. n. sp. (del sur de Baja California Sur). /. (A.) elongata Kevan et al. (Sonora y Sinaloa). /. {A.) costulata
Bolivar (sur de Baja California), /. (.4.) californica Bolfvar (del sur de Baja California Sur). /. (A.) celata. n. sp. (region central del Cabo de Baja
California Sur). y /. (A.) aptera Hebard (de la parte mas meridional de Baja California). El nombre de la ultima especie. Atyphacris californica
(Bruner). se cambia a el nombre presente por ser este un homonimo secundario recurrente. Tablas-claves son provistas. Las relaciones
biogeograficas y filogeneticas son discutidos.

INTRODUCTION

In the course of studying the Pyrgomorphidae of Baja
California, Mexico, preparatory to the publication of a general
handbook on the orthopteroid insects of Baja California that is
currently being coordinated by Dr. D. B. Weissman, I discovered
three hitherto undescribed species of the genus Ichthiacris I.
Bolivar. 1905. Two of these are from the Cape region of Baja
California Sur and the other is from Isla Cedros, off the southwest
coast of Baja California Norte. These and other discoveries now
necessitate a critical re-examination of all species of Ichthiacris.

The Pyrgomorphidae occur primarily in the Old World tropics
and are poorly represented in temperate regions and in the New
World tropics and subtropics. where most of the known species
are Mexican (see Kevan and Akbar 1964. Kevan et al. 1964,
Kevan 1977, 1978). The most northerly range of the American
species is northwestern Mexico (Sonora and Baja California south
of about 29° N latitude), where a single tribe. Ichthiacridini
Kevan, Singh, and Akbar, 1964, occurs.

The Pyrgomorphidae, in general, though not in northwestern

Mexico, are very diverse, but they may be distinguished from all
other Acridoidea by the characteristic form of the male copulatory
structures and the fastigium of the vertex of the head, which has a
distinct median apicodorsal furrow flanked by pair of dorsal
margined depressions, ("areolae" or "foveolae"). Most (and
certainly all American) species also have the upper basal lobe of
the hind femur poorly developed so as not to project forward
beyond the lower lobe, and a rather strongly receding frontal
profile. Neither character, however, is itself diagnostic for the
family. In addition, very many Pyrgomorphidae (including all
New World species except for two of the four known from South
America) are apterous or have extremely abbreviated vestigial
scale-like wings. Again, this is not unique to the family, but, so
far as Mexican genera are concerned, any Acridoidea with more
than mere traces of wings belong to other families.

Three tribes of Pyrgomorphidae occur in Mexico: the
Sphenariini. Ichthyotettigini. and Ichthiacridini. the last having
Ichthiacris as its type genus. The features distinguishing these
tribes are given by Kevan and Akbar (1964) and Kevan et al.
(1964. 1971). In summary, the Sphenariini are robust and

D. K. McE. Kevan

strongly fusiform and (in (he New World) confined to southern
and central Mexico and to Central America: the Ichthyotettigini
are cylindrical, virtually smooth, completely apterous, and found
only in central and southern Mexico; the Ichthiacridini (nymphs
included) are elongate, subfusiform to cylindrical, and have
punctate, granulate, rugose, or somewhat striated integuments,
and most species possess minute vestigial forewings (tegmina).
The male genitalia of the Ichthiacridini. also, are less specialized
than those of the Ichthyotettigini (see Kevan et al. 1964, 1971).
These two tribes overlap to some extent in geographical
distribution, but Ichthiacris is confined to northwestern Mexico
(including Baja California), where the latter tribe is not found.

Prior to the present study, four genera of Ichthiacridini were
recognized (for distinguishing characters, see Kevan et al. 1964):
two from central Mexico. Sphenacris Bolivar. 1884. and
Calamacris Rehn. 1904 (both monotypic). and two from the
northwestern part of the country, the monotypic Atyphacris
Kevan. Singh, and Akbar. 1964. from Baja California, and
Ichthiacris Bolivar. 1905. with three currently recognized species
(Kevan 1978), two from Baja California and one from the
mainland states of Sonora and Sinaloa. Kevan
(1978) — incorrectly, it now proves — synonymized a third Baja
Californian species. /. californica Bolivar, 1905. with /. costulata
Bolivar, 1905 (for which there are several other synonyms), so
that there are really three previously recognized species of
Ichthiacris and one of Atyphacris known from Baja California.

The generic and specific characters and the distribution of the
Ichthiacridini. as previously known, have been outlined by Kevan
and Akbar (1964). and the concealed copulatory structures have
received further attention from Kevan ct al. (1971). A
bibliography for all species, complete to 1976. with the exception
of an overlooked reference by Hebard ( 1923), was published by
Kevan ( 1977). Kevan ( 1978) provided lists of the then-known
localities from which the various species recognized had been
recorded. Inaccuracies in these publications are corrected herein
under the individual species concerned.

MATERIALS AND METHODS

The specimens studied in connection with the present
systematic revision were virtually all dry-pinned and. except for
those in the author's collection, were borrowed in recent years
from various institutions, augmenting the material studied by
Kevan el al. (1964) and Kevan (1978). Preceded by the
abbreviations used for them in the present text, the institutions
involved are as follows: ANSP. Academy of Natural Sciences of
Philadelphia, Philadelphia. Pennsylvania, U.S.A.; CAS,
California Academy of Sciences, San Francisco, California,
U.S.A.; CSLB, Biology Department, California State University,
Long Beach. California. U.S.A.; LEM, Lyman Entomological
Museum. Macdonald College of McGill University, Ste-Anne-de-
Bellevue, Quebec. Canada (which houses the author's collection
of Pyrgomorphidae); SDSNH, San Diego Society of Natural
History. Natural History Museum, San Diego, California. U.S.A.;
USNM. United States National Museum of Natural History,
Washington, D.C.. U.S.A. Unless stated otherwise, material
recorded here for the first time is deposited in CAS or was
handled by, and returned to, that institution.

In the following systematic account, references listed in the

synonymies are limited to primary and otherwise important
citations and to those not included by Kevan (1977). Except for
the new species, details regarding type specimens and their
repositories are given by Kevan et al. (1964) and not repeated
here. Collection data for previously unrecorded material listed
herein are. for accuracy and to facilitate individual recognition,
given virtually as indicated on the labels attached to the
specimens (though the sequence may sometimes be slightly
altered for convenience). They are not converted to a
standardized form as past experience has shown that such
standardization can lead to errors of interpretation. Dates are
expressed in the form "I0.VII.1978." representing, in this case,
"10 July 1978."

The methods of examining the concealed copulatory
structures and the terminology used for these follow Kevan et al.
( 1969). Information regarding habits, life cycles, and autecology
are not discussed herein as this is intended for the handbook
mentioned above.

RESULTS

The genus Ichthiacris is here considered to comprise eight
species. The four species recognized in the earlier revision of
Kevan et al. (1964), despite the inaccuracies in that work, still
stand. These are /. californica Bolivar. 1905. /. costulata Bolivar.
1905, /. elongata Kevan. Singh, and Akbar, 1964, and /. rehni
Bolivar. 1905 (type species); see also Kevan (1977). The
synonymy of /. californica under /. costulata (Kevan. 1978) was
incorrect, and subsequent work has revealed the existence of three
new species — parva. spinifera, and celata — bringing the total to
seven. /. celata. however, possesses certain characters (notably
the virtual or complete absence of tegminal vestiges and the form
of the phallic structures) that now make it impossible to regard
the hitherto monotypic Atyphacris as generically distinct from
Ichthiacris. This, in turn has resulted in the recurrence of
secondary homonymy. The type species of Atyphacris is A.
californica (Bruner, 1906), which, on being transferred back to
Ichthiacris. again becomes a junior homonym of /. californica
Bolivar, 1905, so that it must revert to its previous replacement
name. /. aptera Hebard. 1932, and is the eighth species of that
genus.

The type species of Ichthiacris. I. rehni, is now seen to stand
apart, more so than does /. aptera, from other previously
described species of the genus, notably in its more rugose
integument, generally broader tegminal vestiges (particularly in
the female), deeper dorsal ovipositor valves, and the form of the
phallic structures. Prior to the present revision, it would have
indeed been plausible to transfer all other previously known
species from Ichthiacris to Atyphacris and to let the latter remain
a full genus. Nevertheless, though one of the newly discovered
species, close to /. rehni. namely. /. parva. would support such an
action, another, /. spinifera. possesses certain characters
indicating relationship to /. rehni but others (narrower tegminal
vestiges, shallower dorsal ovipositor valves, and the form of the
aedeagus) more in keeping with those of the remaining species.
Therefore I have adopted a compromise by recognizing two
subgenera: Ichthiacris. sensu stricto, for/, rehni and the two new
species just mentioned, and /. {Atyphacris) for the remaining five
(including one new) species. With the exception of /. (A.)

The Genus Ichthiacris

elongata, from the northwestern Mexican mainland, all species of
Ichthiacris are confined to the southern half of Baja California.

SYSTEMATICS

GENUS ICHTHIACRIS I. BOLIVAR, 1905. SENSU LATO

Ichthiacris Bolivar. 1905: 287; Kevan. Singh, and Akbar. 1964: 234, 239.

Calamacris (nee Rehn); Bruner. 1906: 200.

Atyphacris Kevan. Singh, and Akbar, 1964: 233, 240; syn. nov. (now

subgenus only).

Type species. — By subsequent designation (Kirby 1910),
Ichthiacris rehni Bolivar. 1905.

Diagnosis. — Body elongate-fusiform to subcylindrical,
usually rather slender, with distinctly punctate, granulate,
pustulate, or striated integument; almost (sometimes completely)
apterous; copulatory structures relatively unspecialized for the
Pyrgomorphidae (see Kevan et al., 1964. 1971 ).

Remarks. — The combination of characters noted readily
distinguishes the Ichthiacridini from all other American
Pyrgomorphidae. The monotypic genera Sphenacris and
Calamacris differ from Ichthiacris in possessing more definitely
elongate-fusiform bodies and more strongly pustulate
integuments. There are also important genitalic differences, as
indicated by Kevan et al. (1964. 1971). Sphenacris and
Calamacris appear to be restricted to central Mexico.

Two subgenera of Ichthiacris are here recognized,
distinguishable from each other by means of Table 1 .

Subgenus Ichthiacris Bolivar. 1905, sensu stricto

Ichthiacris Bolivar, 1905: 287 (partim); here restricted.

Type species. — As for genus.

Diagnosis. — Sculpture strongly rugosostriate. usually with
some tubercles: fastigium of vertex always considerably longer
than basal width (Figure 1A-D, N, O); frontal profile strongly
oblique, rather strongly and sinuously concave (Figure 2A-D. N.
O); inferoposterior region of lateral pronotal lobe typically rugose
(Figure 3A, B, I), angle sometimes with a small spine or sharp
denticle (as in Figures 2C, 3B); minute tegminal vestiges present,
sometimes slip-like, typically broad and lobe-like (as in Figure
3A).

Males: Abdominal terminalia dorsally as in Figure 4A, B, H;
subgenital plate in lateral view fairly acute (Figure 5A. B, H);
epiphallus with posterior emargination broad or shallowly U-
shaped (but not deeply so nor slot-like), anterolaterally directed
subterminal processes of lateral appendices prominent (Figure
6A-C, O): apical parts of aedeagal valves and sclerites relatively
short (Figure 7 A, B, O) or of moderate length (Figure 7C).

Females: Metathoracic epimera not expanded dorsally (Figure
3A, B. I), sometimes with tubercles (Figure 3A. 1): dorsal
ovipositor valves in lateral view typically deep and apically
sickle-like (Figure 8A) but sometimes tapered (Figure 8B);
subgenital plate dorsally with posterior margin on either side of
egg-guide having short, distinct, closely set, longitudinal
striations extending about half-way along margin (Figure 9A. B.
H); receptacula seminis as in Figure 10A, B.

Distribution. — Baja California, Mexico, only (Figure 1 1 ).

Included species. — /. (/.) rehni Boh'var, 1905; /. (/.) parva, n.

sp. (Figure 12), and /. (/.) spinifera, n. sp. (Figures 13A, B, 14A,
B). They are separable from each other as indicated in Table 2.

Ichthiacris (Ichthiacris) rehni, Bolivar, 1905 (Figures 1A, B; 2A.
B; 3A; 4A; 5A; 6A. B; 7A. B; 8A; 9A; 10A; 1 1 1

Ichthiacris rehni Bolivar, 1905: 287, 288.

Ichthiacris rehni; Kevan, Singh, and Akbar. 1964: 236. fig. 1

(map — partim), 240, 241. fig. 4 (parrim), 242 (partim), 243, fig. 5,

288. 289 (plate 1), figs. 1-L; Kevan, 1977: 65, 665 (partim— by

inference, referring to foregoing), 644; Kevan, 1978: 5, fig. 1

(map — partim), 15, 23-24 (partim); Kevan, 1981: (28).

Recognition. — Integument rugose, small spine or denticle on

the inferoposterior angle of the lateral pronotal lobe lacking;

frontal profile strongly concave and sinuous in both sexes (Figure

2A, B); tegminal vestiges (Figure 3A) broad (often as broad as

long in female: at least half this in male); dorsal ovipositor valves

deep and apically sickle-like (Figure 3A). Other characters are

noted in Table 2.

Distribution (Figure 11). — The type specimens (including
male lectotype) were described merely from "Basse Californie."
The species is quite widely distributed in southern Baja California
from a little north of 28°N almost to 23'30'N latitude. In addition
to the types, the specimens studied are listed below. Specimens
noted in earlier publications are indicated only by locality and
reference; those recorded erroneously as this species are omitted
here but are listed under the appropriate species.

Material examined. — Baja California Norte: 7 km N of
Guerrero Negro [itself just within Baja California Sur] at 0.24 km
N of km 121 on Mex[ico Highway] 1. Stop 34, 10.VII.1978, D.
B. Weissman and D. C. Lightfoot, 2 juvs. (6 9 ); 68 [road] km S
of [Rancho] Rosarito, 0.2 km N of km 121 on Mex[ico Highway]
1 [i.e., between Rancho Mezquital and Colonfa Agricola].
25.VII.1977. [no collector]. 1 juv. (6).

Baja California Sur: 31 .0 km NW of Punta Abreojos, elev. 62
m, sand flats. Encelia halimifolia Cav.. 21.1.1975. E. L. Sleeper.
2649/11341. 2 9 9 (CSLB); Arroyo Seco (Kevan et al. 1964,
Kevan 1978); 7 rd. mi. NW, 1.1. mi. SW of El Cien [ca. 24°22'N.
110°58"W], on Santa Fe road, elev. 100 feet. 13.IX.1983. J. P. and
K. E. Donahue, 1 9 ; 46 km W of Col[onia] Constitucion,
19.X. 1974, D. Otte, 16 6,299 (ANSP except 2 66, 1 2
LEM); Loreto (Kevan et al. 1964. Kevan 1978) [adult 9; large 9
juvs. with same data recorded by Hebard (1923); see /. (A.)
costulata]; Mangrove I., Magdalena Bay (dunes), 13. III. 1966. C.
L. Hogue, 3 6 6 (1 LEM); La Paz (Kevan 1978); 4-9 and
14-22.IX.1967. J. Chemsak and A. L. M. Michelbacher. 2 juvs,
(99); elev. 25 feet. 30.IV1969, S. C. Williams. 1 9; NW of La
Paz and 59 mi. NW of La Paz (Kevan 1978) (latter 9. not 6 ); 70
mi. N of La Paz, 19. VII. 1974, D. Otte, 1 9 (ANSP); 73 km N of
La Paz, elev. 320 m, 19.X.1974, M. Descamps, 1 6, 1 2 (ANSP);
12 mi. N of La Paz on road to Pichilingue (Kevan et al. 1964.
Kevan 1978): 5 mi. SW of La Paz. 7. IX. 1967, [no collector,] 1 9;
18 mi. W of La Paz, 7.X.1983. F. Andrews and D. K. Faulkner. 1
9 (SDSNH); ca. 44 km W of La Paz at 0.2 km S of km 44 on
Mex. Hwy. 1, Stop 79-14, 31. XII. 1978. D. B. Weissman et al., 1
(5,1?; 43.5 km W of La Paz at km 43.5 on Mex. Hwy. 1, Stop
79-30. 4.1.1978, D. B. Weissman et al., 1 9; 3.2 km S. [Rancho]
Las Pocitas at km 109 on Mex. Hwy. 1, elev. 60 m, Stop 79-21 1,
28. IX. 1979, D. B. Weissman et al., I 6, 2 9 9. 1 juv. ( 9 ); 3 mi.

D. K. McE. Kevan

Table 1. Characters distinguishing subgenera of lchthiacris.

Character

/( hthiacris sensu stricto"

Atyphacris"

Known distribution
Sculpture

Head
Fastigium of vertex (dorsal I
Vertex proper (lateral view)

Frontal profde
Lateral pronotal lobe
Inferoposterior region

Metathoracic epimera

Tegminal vestiges

Males
Epiphallus
Posterior emargination

Anterolateral!;, directed
subterminal processes
of lateral appendices

Endophallus

Apical parts of aedeagal
valves and sclerites

Females

Ovipositor

Dorsal valves in profile

Subgenital plate (dorsal)

Posterior margin on either
side of egg-guide

Baja California only

Strongly rugosostriate. with some
(sometimes rather acute) tubercles'

Always distinctly longer than basal width

Abruptly convex above eyes, at
least in males

Strongly concave, usually sinuously

Often rugose, angle sometimes
with small spine or denticle'

Not expanded, sometimes
tuberculate

Present, sometimes slip-like but often
lobe-like.' particularly in females

Broadly excavated to shallowly
U-shaped

Prominent to exaggerated

Rather short or length moderate

Deep and apically sickle-like,'
or tapered

With distinct, short, longitudinal
stnations extending about
hall-way along margin

Baja California. Sonora. Sinaloa

Not so strongly striate.'' somewhat rugose, granular or
punctate; tubercles feeble or lacking'

Variable, sometimes little if at all longer than basal width
Evenly or only weakly convex

Usually only weakly concave, seldom sinuously

Seldom rugose, angle without
spine or denticle

Sometimes expanded in females,
not tuberculate

Always narrow and slip-like
or absent

Deeply U-shaped or slot-like

Not prominent

Relatively short to very long

Always tapered, not deep

With indistinct or weak crenulations
only, extending for but a short
distance along margin

"Figures 1 A-D, N, O, 2A-D, N. O. 3A. B, I. 4A. B, H, 5A, B. H, 6A-C. O, 7A-C,
'•Figures 1E-M. 2E-M, 3C-H, 4C-G. 5C-G, 6D-N, 7D-N. 8C-E, 9C-H, 10C-H.
' More obvious external feature.
''Young nymphs are often distinctly more striated than adults.

O. 8A. B, F, 9A. B, I. 10A, B. 11, 12. 13A, B, 14A. B.
13C. 14C. 15, 16.

The Genus Ichthiai n's

ATA

D. K. McE. Kevan, 1990. Proc. San Diego Soc. Nat. Hist. 1: 1-34.

Page 5, caption to Figure 1:

end of line 2: for "holotype; 0, I." read "?; N, I."

line 3: for "?; N," read "*, holotype; 0," and delete "<f,"

ERRATUM

5 mm

M

Figure 1. Dorsal view of heads of Ichthiacris spp. A. B. /. (/.) rehni, 5. 9; C, I. (I.) spinifera, 6, holotype; D, /. (1.) spinifera, 9 allotype; E. F, /.
{Atyphacris) elongata, 3. 9; G, H. /. {A.) costulata, 6. 9: 1. J. /. M.) californica, 6. 9: K. /. [A.) celata, i: L, M, /. (A.) aptera, 6, holotype; O. /.
(I.) parva, 9; N. /. [I.)parva, 6", 9 (last-instar juv.).

D. K. McE. Kevan

Table 1. Characters distinguishing subgenera of Ichthiacris.

Character

Ichthiacris sensu stricto"

Known distribution
Sculpture

Head
Fastigium of vertex (dorsal)
Vertex proper (lateral view I

Baja California only

Strongly rugosostriate, with some
(sometimes rather acute) tubercles'

Always distinctly longer than basal width

Abruptly convex above eyes, at
least in males

AtyphacrisP

Baja California, Sonora. Sinaloa

Not so strongly striate,1' somewhat rugose, granular or
punctate; tubercles feeble or lacking'

Variable, sometimes little if at all longer than basal width
Evenly or only weakly convex

ai'i'Mn Giniimielv

Dorsal valves in profile

Subgenital plate (dorsal)

Posterior margin on cither
side of egg-guide

Deep and apically sickle-like.'
or tapered

With distinct, short, longitudinal
striations extending about
hall -way along margin

Always tapered, not deep

With indistinct or weak crenulations
only, extending for hut a short
distance along margin

"Figures 1 A-D. N. O. 2A-D. N. O. 3A. B. I. 4A. B, H, 5A, B, H. 6A-C. O, 7A-C, O. 8A. B. F, 9A, B. I, 10A. B, 1 1, 12, 13A, B, 14A, B.
''Figures 1E-M, 2E-M, 3C-H. 4C-G. 5C-G. 6D-N, 7D-N. 8C-E. 9C-H. 10C-H, 13C, 14C, I?. 16.

' More obvious external feature.

''Young nymphs are often distinctly more striated than adults.

The Clonus Ichthiacris

Figure I. Dorsal view of heads of Ichthiacris spp. A, B. /. (/.) rehni, 6. 9; C. I. (I.) spinifera, 6, holotype; D. / (/ I spinifera, 2 allotype; E, F. /.
(Atyphacris) elongata, 6, 2; G. H, /. (A.) costulata, 6. 2; I, J, /.(A.) californica, i . 2; K.I. (A) celata, 6: L. M. /. (A.) aptera, 3. holotype; 0, /.
(I.)parva, 9 ; N, /.(/.) parva, 6. 2 (last-instarjuv.).

D. K. McE. Kevan

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D. K. McE Kevan

5 mm

H

Figure 4. Dorsal view of abdominal terminalia of 6 Ichthiacris spp. A. /. (/.) rehni
1 / i caIifornica;F,[.(A.) celata:G,I.(A.) aptera;H,l.(I.) parva.

B, /. (/.) spinifera; C, /. (Atyphacris) elongate; D. /. {A.) costulata;

The Genus Ichthiacris

5 mm

Figure 5. Lateral view of abdominal terminalia of 8 Ichthiacris spp. A, /. (/.) rehni; B, /. (/.) spinifera (the angle of viewing makes the subgenital plate
appear proportionately shorter than it really is); C, /. {Atyphacris) elongata; D, /. {A.) costulata; E. /. (4.) californica; F, /. iA.) reluia: G. /. (.4.)
aptera; H, /. (I.) parva.

10

D. K. McE. Kevan

Irnm

Figure 6. Epiphalli of 6 Ichthiacris spp. A. B. /. (/.) rehni (A, 44 km W of La Paz.; B. Arroyo Seco); C, /. (/.) spinifera, holotype; D, /. (Atyphacris)
elongata, paratype, CAS; E-H, /. {A.) costulata (E. El Arco; F. road to La Burrera; G. 10 km S of San Pedro; H. 44 km W of La Paz); I-L. /. (A.)
californica ( I, El Triunfo; J. 4.3 km SE of Valle Perdido; K, La Burrera; L. 1 1 .2 km S of Todos Santos); M, /. (A.) celata, holotype; N. /. {A. ) aptera,
paralectotype; O. /. (/.) parva, paratype. LEM KPY-21-05-P4.

The Genus Ichthiacris

II

Figure 7. Aedeagi of 6 Ichthiacris spp. . A, B, /. (/.) rehni (A, 44 km W of La Paz; B, Arroyo Seco); C. /. (/.) spinifera, holotype; D. /. {Atyphacris)
elongata, paratype, CAS; E-H, /. (A.) costulata (E, El Arco; F, Puerto Escondido; G, road to La Burrera; H. 44 km W of La Paz); I-L, /. {A.)
californica (I, El Triunfo; J, 4.3 km SE of Valle Perdido; K, La Burrera; L, 1 1.2 km S of Todos Sanlos); M. /. (A.) celala, holotype; N. /. (-4.) tiptera,
paralectotype; O, /. U.)pana. paratype. LEM KPY-21-05-P4.

12

D. K. McE. Kevan

5 mm

Figure 8. Laleral view of abdominal terminalia of 5 Ichthiacris spp.. showing ovipositor. A. /. (/.) rehni; B, /. (/.) spinifera; C. /. (Atyphacris) elongata;
D, /. 04.) i ostulata \l. (.4. 1 californit a similar]; E. /. [A.) aptera; F. /. (I.)parva dasi-insiar juv.).

The Genus Ichthiacris

13

Figure 9. Dorsal view of subgenital plates of 9 Ichthiacris spp. A, /. (/.) rehni (43.5 km W of La Paz); B. /. (/.) spinifera, paratype. LEM KPY-21-06-
P3; C, /. (Atyphacris) elongata, paratype, LEM KPY-21-03-P3: D. E, /. (A.) coslulata (D, Puerto Escondido; E, 25 km S of San Ignacio): F. G, /. (A.)
californica (F, La Bun-era; G, El Triunfo); H, /. (A.) aprera (La Burrera): I. /. {!.)pana. last-instar juv. (allotype)

14

D. K. McE. Kevan

Figure 10. Receptacula seminis of ? Ichthiacris spp. A, /. (/.) rehni (43.5 km W of La Paz); B, /. (/.) spinifera, paratype. LEM KPY-21-06-P3; C. /.
(Atyphacris) elongara, paratype, LEM KPY-2I-03-P3; D, E, /. (A.) costulata (same spec-imens as in Figure 9); F. G. /. (A.) californica (same specimens
as in Figure 9); H, /. [A.) aptera (La Burrera).

The Genus Ichthiac

15

Figure 1 1. Known distribution of Ichthiacris, sensu stricto. Circles, /. (/.) rehni: squares, /. (I.)pana; triangles, /. (/.) spimfera.

16

D. K. McE. Kevan

cm

&

Figure 12. Ichthiacris (l.)parva, n. sp., type specimens. A. 8 holotype, dorsal view. B. (he same, lateral view; C, last-instar 9 nymph, allolype. dorsal
view; D, the same, lateral view.

The Genus Ichthiacris

17

1 cm

Figure 13. Ichthiacris n. spp.. type specimens, dorsal view. A, /. (/.) spimfera. 6 holotype; B, the same, 9 allotype; C. /. {Atyphacris) celata, 6
holotype.

18

D. K. McE. Kevan

Table 2. Characters distinguishing species of Ichthiacris sensu stricto.

Character

1 .(/ '.) rehni"

I (I )pan-ah

I. (/.) spinifera1

Known distribution
Size (mm body length)
Sculpture (striation)
Frontal profile

Inferopostenor region of lateral
pronotal lobe

Tegminal vestiges

Males

10th abdominal tergum (dorsal)

Posterior dorsolateral
projections

Cerci

Epiphallus

Anterolaterally directed
subterminal processes of lateral
appendices

Endophallus

Apical in relation to basal pairs
of aedeagal valves

Females

Ovipositor

Dorsal valves in profile

Baja California from a little N of Isla Cedros. SW Baja California

28°N to almost 23 1/2°N Norte, only

Averaging larger: d 18.5-25; Very small: d 14.5-15.8; 9 < 25^
926.5-45

Strong, usually with prominent Moderately strong, with pustular

irregularly arranged tubercles'' tubercles (Fig. 12)

Sinuous in both sexes (Fig. 2A. B) Sinuous in both sexes (Fig. 2N. O)

Usually without prominent granules Usually with indications of

or tubercles; angle without spine or tubercles"; angle without spine or

denticle (Figure 3A). denticle (Fig. 2N, O)

Generally broad (often as broad as Generally broad in females (so far

long) in females; usually at least half as known): narrow and scale-like

as broad as long in males (Fig. 3A) in males (Figs. 2N and 31)

Pointed, sometimes inwardly
directed (Fig. 4A)

Relatively robust, little incurved
(Figs. 4A and 5A)

Relatively large (Fig. 6A, B)

Very large and prominent
(Fig. 6A. B)

Relatively large (Fig. 7A. B)
Shorter (Fig. 7 A. B)

Broadly triangular (Fig. 4H)

Relatively robust, little incurved
(Figs. 4H and 5H)

Small (Fig. 60)

Small, acute (Fig. 60)

Small (Fig. 70)
Shorter (Fig. 70)

Baja California Sur S of 24°N
only

Moderate: d 18.5-23; $ 29-33

Less strong, with few tubercles
(Figs. 13 and 14A. B)

Sinuous in males, less or scarcely
so in females (Fig. 2C. D)

Usually with small, granular
tubercles; with small spine or
denticle (Figs. 2C and 3B)''

Narrow and slip-like in both sexes,
less than half as broad as long
(Figs. 2C and 3B)

Rather broadly triangular (Fig. 4B)

Slender, distinctly incurved
(Figs. 4B and 5B)

Moderate-sized (Fig. 6C)

Moderate-sized (Fig. 6C)

Moderate-sized (Fig. 7C)
At least as long (Fig. 7C)

Deep, apically sickle-like (Fig. 8A)'' Apparently rather deep (known

only for last-instar nymph. Fig. 8F)

Subgenital plate (dorsal)

Sclerotized areas on either side Transversely elongate-triangular
of egg-guide (Fig. 9A)

|Unknown (nymph. Fig. 91)

Relatively more slender and
tapering, not apically sickle-like
(Fig. m)d

Irregularly subquadrate (Fig. 9B)

"Figures 1A. B. 2A. B, 3A, 4A, 5A, 6A. B. 7A, B, 8A. 9A, 10A. II.
^Figures IN, O, 2N, O, 31, 4H, 5H, 60. 70. 8F. 91, 11. 12.
'Figures 1C, D, 2C. D. 3B, 4B, 5B, 6C, 7C. 8B, 9B, 10B, 11, I3A, B, 14A, B.
''More obvious external feature

The Genus Ichthiacris

19

SE of Pozo Grande (Kevan 1978); Isla Magdalena, Howland's
Lagoon, sand dunes. Stop 83-93. 8.VII.1983. D. B. Weissman and
V. F. Lee, 8 8 8 (mostly rather small). 13 2 2 (generally rather
small to very small), 1 8 juv. (?2nd instar), 1 2 juv. (last instar)
(3 8 8, 4 9 9 LEM); Isla Magdalena, canyon 1 km NW of Puerto
[Bahfa] Magdalena, Stop 83-94, 8. VII. 1983, (D. B. Weissman],
D. K. Faulkner and D. C. Lightfoot, 1 2 (rather small) (SDSNH);
[Puerto] San Carlos. Magdalena Bay, 25.IX.1981. D. Faulkner. 1
d, 1 2 (SDSNH); 12 mi. S of La Purisima [as "Colonia
Puerissima"). 5.X.1967. G. A. Marsh. 1 2 (very large) (LEM); 19
mi. SW of San Miguel Comondii (Kevan 1978); Isla Santa
Margarita. 3 km SW of Puerto Cortes, sand dunes. Stop 83-92.
7.VII.1983, D. B. Weissman and D. C. Lightfoot, 3 8 8 (rather
small), 2 2 2 (rather small), 2 2juvs. (1 last instar; 1 small, ?2nd
instar); Santa [as "San"] Rita (Kevan 1978); 2 mi. SE of Santa
Rita (Kevan 1978); 13.5 km N of Todos Santos, elev. 46 m. Cape
thorn forest, beaten from Hymenoclea monogyra Torr. and Gray.
7-8.1.1981. E. L. Sleeper, 2333/10948, 1 3,1 8 juv. (last instar)
(CSLB); [Ejido] Vizcaino. 69.5 km SE of Guerrero Negro, elev.
121 m, wash, on Hxmenocle monogyra, 22. XL 1980. E. L.
Sleeper, 2742/1 1325, 1 2 (CSLB); 21.6 km W of Ejido Vizcaino,
elev. 90 m. disturbed sand dune, Encelia sp.. taken at 1600 hrs.
11.1.1980. E. L. Sleeper, 2740/11340. 1 8 (CSLB); 68.0 km W of
Ejido Vizcaino, elev. 30 m. high stable sand dune with typical
climax vegetation, Encelia laciniata Vasey and Rose,
11-12.8.1989, E.L. Sleeper, 2730/11355. 2 8 8,2 99 (CSLB).

Remarks. — Because of a paucity of specimens in collections,
there has been some confusion in the literature between this and
other species of Ichthiacris. The female specimens recorded from
"Mesquital" [presumably Rancho Mezquital in the extreme south
of Baja California Norte — but see distribution of/. (A.)
costulata], from 20 mi. S of El Arco. from Loreto. and from [El|
Triunfo by Kevan et at. (1964) and repeated by Kevan (1978)
belong, in fact, to /. (A.) costulata (and from the last two localities
the specimens are last-instar nymphs and not adult females!). The
same would seem to be the case also with the nymphs recorded as
/. rehni by Kevan et ai. (1964) from the Sierra de San Lazaro
[now Sierra de la Victoria], as already noticed by Kevan (1978),
from the islands of Coronados. Danzante. and San Marcos, and
from 1.3 mi. NW of El Triunfo (all repeated, uncorrected, by
Kevan 1978). The nymph recorded from [2.7 mi. SE of] Valle
Perdido by Kevan (1978) now proves to belong to the new
species /. (/.) spinifera.

The inclusion by Kevan et al. (1964) of the females
mentioned above under /. rehni was unfortunate as it resulted in
the misrepresentation of this species in some of the line
illustrations in their figure 4 (p. 241). The lateral view of the
female's head should be more like that of the male's, in which the
sinuous outline of the frontal profile is more strongly indicated.
In addition, the ovipositor valves in the lateral view are shown as
being tapered, whereas, in /. (/.) rehni, they are deeper and
apically sickle-shaped. The lateral views of the meso- and
metathoracic regions, alleged to be of/. (/.) rehni by Kevan et al.
1964. fig. 4E), are quite typical for some females of /. (A.)
costulata, not for /. (/.) rehni. This is because the metathoracic
epimeron (narrower than in most specimens of the former
species) is distinctly convex instead of being rather straight
dorsally. In /. (A.) costulata the metathoracic epimeron is quite
variable in shape, usually being intermediate between the

condition indicated by Kevan et al. ( 1964: 241. fig. 4E). allegedly
for /. (/.) rehni, and thai illustrated by them for /. (A.) costulata
(their p. 246. fig. 6E). Kevan (1978) noted that the metathoracic
epimera of /. costulata are wider and more "flared" than in /.
rehni. but he did not comment on the erroneous nature of the
figure referred to above. With regard to the shape of the tegminal
vestiges, Kevan (1978) stated that a narrow form is unusual in /
rehni. In fact, the tegminal vestiges of female /. rehni are less
than twice as long as wide and never as narrow as in the
illustration of Kevan et al. (1964: 241, fig. 4E). Even in males
they are at least a little wider than shown. In referring to their
illustration of a female paralectotype of /. rehni, Kevan et al.
(1964: 288. caption to plate I) also suggested that its strongly
rugosotuberculate integument was the less common condition of
the species. Later. Kevan (1978) indicated the reverse to be the
case. All less rugose specimens having narrow slip-like tegminal
vestiges have proved to belong to /. (A.) costulata.

Ichthiacris (Ichthiacris) parva, new species
(Figures IN, O; 2N, O; 31; 4H; 5H; 60; 70; 8F; 91: 11

12)

Holotype. — Figure 12 A, B. CAS, type no. 15710. Mexico:
Baja California Norte. Isla Cedros. Bahia del Sur, "Playa Playon."
28. IX. 1984. Leafless shrubs bordering sandy dunes ca. 100 m
from ocean; at night. D. B. Weissman and V. F. Lee. Stop 84-63.
8.

Allotype. — Figure 12C, D. CAS. type no. 15710, Mexico:
same data as holotype. Last-instar nymphal 2 .

Paratopes.— CAS, 2 8 8; LEM, type no. KPY-21-05-P4, Id,
Mexico: same data as holotype.

Recognition. — Similar in most features to /. (/.) rehni, but size
much smaller (cT < 16, 2 < 25 mm) and phallic structures
differing as indicated (compare Figures 60 and 70 with Figures
6A, B and 7A, B). Other characters as indicated in Table 2.

Etymology. — Small, referring to the diminutive size.

Description of holotype. — Body moderately elongate and
rather slender, coarsely rugosostriate throughout, with some
scattered pustular tubercles on head and thorax; hairs of frons.
sternum, legs, and end of abdomen neither long nor dense.

Antennae about as long as head and pronotum together, with
12 articles in addition to scape and pedicel, terminal article
conical, about three times as long as wide, other articles, except
scape, subquadrate; articulation between two basal articles of
flagellum indistinct.

Head (Figures IN and 2N) twice as long as basal width, a
little longer than pronotum; eyes large and prominent, elliptical,
depth about two-thirds of length, interocular space at narrowest
about half greatest dorsal width of an eye; fastigium of vertex
bluntly elongate-triangular with slightly sinuous margins, about
1.75 times as long as basal width, with a pair of strong, rather
irregular, mediolateral longitudinal ridges: median sulcus between
dorsal areolae extending backward for nearly one-third of length
of fastigium before giving way to a distinct medial cannula that
extends backward as a strong, raised ridge between eyes,
becoming obsolescent in the occipital region; vertex proper
abruptly convex in lateral view; eyes margined dorsally by rather
strong oblique ridges; frontal profile very strongly oblique,
concave and somewhat sinuous; fastigium of frons almost straight
below and nearly parallel to dorsum of fastigium of vertex, which

20

D. K. McE. Kevan

it meets apically at a moderately acute angle (as in Figure 2N);
frontal ridge very prominent above antennal bases, strongly and
narrowly sulcate below it to the median ocellus, beneath which it
is less prominent as far as clypeus; lateral frontal carinae. in
lateral view, strong and sinuous, passing close to ventral margins
of eyes, in frontal view also moderately sinuous.

Thorax (Figure 2N): Pronotum subcylindrical; disk with
median carina well defined in front of, but weaker behind, median
transverse sulcus; lateral carinae virtually obsolete, anterior
margin subtruncate. posterior margin weakly biarcuate; posterior
transverse sulcus almost straight, crossing pronotal disk at about
four-fifths of pronotal length; median transverse sulcus also
almost straight, crossing disk distinctly behind its middle; lateral
pronotal lobe considerably longer than deep, its anterior margin
oblique, almost straight, its inferior margin strongly sinuous and
its posterior margin almost vertical but concave and with
prominently protruding ends to longitudinal ridges of lobe;
inferoposterior angle obliquely truncate and with a rugose area
immediately anterior to it. lacking a spine or denticle (Figure 2N).
Mesonotum transverse, distinctly shorter than either metazona or
mesozona (region between median and posterior sulci) of
pronotum. Metanotum about 1.5 times as wide as long;
metathoracic epimera very narrow; prosternal tubercle stout,
broadly conical, rounded apically. flattened anteriorly;
mesosternal lobes almost twice as long as wide, their interspace
considerably wider than a lobe.

Wings: Tegminal vestiges (Figure 2N) very small, parallel-
sided, blunt apically, extending only slightly beyond posterior
margin of mesonotum; hindwing vestiges (if any) not visible.

Legs: Front femora barely as long as pronotum. weakly
incrassate and feebly carinate dorsally; middle femora similar to
front femora, but more flattened than incrassate; hind femora
moderately slender, about 4.6 times as long as greatest width, at
rest not quite reaching apex of abdomen.

Abdomen: Median dorsal carina on terga I— VII strong, well
defined, that on tergum I exaggerated toward posterior margin;
tympana lacking; terminalia as in Figures 4H and 5H; tergum X
with posterior emargination shallow, of moderate width, concave;
dorsolateral processes bluntly triangular; epiproct broadly
triangular, not much longer than basal width; cerci comparatively
short and stout, barely reaching apex of epiproct; subgenital plate
acute in lateral view.

Phallic structures (from LEM paratype, not extracted from
holotype; Figures 60, 70): Epiphallus small with comparatively
narrow bridge; lateral plates broadly expanded basally; posterior
margin broadly and shallowly U-shaped; lophi short and
dorsolateral^ directed: lateral appendices stout, of characteristic,
irregular form with subterminal processes prominent, pointed, and
much smaller than in other species of subgenus Ichthiacris
(compare Figure 60 with Figures 6A-C); endophallus small with
apical parts of aedeagal sclerites and valves short and blade-like
(Figure 70). much as in /. (/.| rehni (Figure 7A. B).

Coloration grayish, body mottled fuscous with many blackish
pustular tubercles; cheeks margined by blackish above; inferior
regions of lateral pronotal lobes and of genicular lobes of hind
femora all somewhat paler; eyes pale brown with six curved
blackish stripes; tegminal vestiges with dorsal (posterior)
margins reddish; hind femora with carinae of outer faces flecked
with elongate blackish maculations, inner faces rather uniform

grayish; hind tibiae blackish-gray above.

Measurements (mm): Length of body 15.8: antenna 5.0: head
3.0; pronotum (mid-dorsal) 2.1; tegmen vestige 0.65; hind femur
6.9; maximum width of hind femur 1 .4.

Description of Allotype. — [No adult female is available, but in
this genus last-instar nymphs resemble adults sufficiently closely
for valid description.] The specimen agrees in general with the
description of the holotype but differs in its slightly larger size, in
its abdominal terminalia, and in the following:

Antennae more ensiform, considerably shorter than head and
pronotum together, with short, strongly transverse articles. [In an
adult female, the antennae would have slightly longer articles than
in this allotype.]

Head (Figures lO and 20) relatively shorter, very little longer
than pronotum; eyes less prominent, greatest depth less than two-
thirds of length; interocular space at narrowest about as wide as
greatest dorsal width of an eye [in an adult female the space
would doubtless be a little wider still]; vertex proper, in lateral
view, not so strongly convex between eyes; frontal profile a little
less oblique, not quite so sinuous.

Thorax (Figure 31): Pronotum, in dorsal view, slightly
divergent posteriorly, lateral carinae indicated in prozona. lateral
lobe with inferior margin nearly straight, posterior margin less
concave and inferoposterior angle less truncate than in holotype;
mesonotum a little less transverse and prosternal tubercle a little
more transverse than in holotype; mesosternal lobes not quite so
long as wide, their interspace with concave margins, least width
of latter greater than the greatest width of a mesosternal lobe.

Wings: Buds of tegminal vestiges (Figure 31) broad and lobe-
like (that of left being wider and more oval than that of right), not
extending beyond posterior margin of mesonotum. [In an adult
female the tegminal vestiges would be similar but perhaps a little
larger in proportion.]

Legs: Front femora shorter than pronotum and not incrassate.

Abdomen: Medial dorsal carina not exaggerated on tergum I;
terminalia as in Figure 8F; epiproct a little broader and blunter
than in male; cerci short, stout, conical, not half so long as
epiproct; ovipositor (Figure 8F) with dorsal valves moderately
deep in lateral view but smooth, in accord with nymphal status;
subgenital plate in dorsal view (Figure 91) with egg-guide short
and bluntly triangular [ in an adult female it would doubtless be a
little longer and more acute], posterior margin on either side with
short longitudinal striae sparse but extending for about half width
of margin [in an adult female they would presumably be denser],
columellae and associated sclerotized patches undeveloped:
spermatheca lacking, in accord with nymphal status.

Coloration same as that of holotype.

Measurements (mm): Length of body 18.2; antennae ca. 4
(curved); head 3.8; pronotum 2.8; tegminal vestige bud 0.9; hind
femur 7.0; maximum width of hind femur 1.5. [An adult female
would presumably not exceed 25 mm in body length.]

Paratypes. — These agree quite closely with the holotype,
though two (CAS) are even smaller, the smaller of these being
only 14.5 mm in body length; another (LEM) has tegminal
vestiges proportionately even smaller than those of the holotype,
though in all specimens these are distinctly longer than wide. The
coloration does not vary significantly, though it is a little darker
(presumably because of postmortem discoloration) in two
specimens.

The Genus Ichthiacris

21

Distribution (Figure II). — Known only by the type series
from Isla Cedros, off the southwestern coast of Baja California
Norte.

Cytology. — I am informed by D. B. Weissman (in litt., 1984)
that the karyotype of this new species differs from that of other
species of Ichthiacrts examined by him in that, although 2nd =
19, as is usual for Pyrgomorphidae, "there is only 1 small
chromosome pair versus the normal 2; there are the normal 2
large bivalents during meiotic metaphase I. but [there are] 6
medium-sized bivalents versus the usual 5." Two specimens in
anaphase II were checked to rule out the possibility of a
heterozygous supernumerary segment.

Remarks. — This species would seem to be an insular
derivative of the more widely distributed /. (/. ) rehni and to have
evolved in isolation. The epiphallus is anomalous, exhibiting, in
comparison with other species of Ichthiacris. certain more
primitive features. These include the divergent lophi
characteristic of the subgenus Atyphacris, the more U-shaped
posterior margin characteristic of /. (A.) californica [but with
basally widened lateral plates, as in some /. (A.) costulata], and a
curious, derived form of the lateral appendices (accompanied by a
reduction in the size of their subterminal processes).

Ichthiacris {Ichthiacris) spinifera, new species

(Figures 1C, D; 2C, D; 3B; 4B; 5B; 6C; 7C; 8B; 9B; 10B; 13A,

B: 14A. B)

Ichthiacris rehni (nee Bolivar): Kevan, 1978: 23 (partim — 1 juv., [2.7 mi.

SE[ Valle Perdido. only).

Holotype.— Figures 13A, 14A. LEM, type no. KPY-21-06-H,
Mexico: Baja California Sur], Arroyo [de] San Bartolo, 1 mi. SE
San Bartolo, 20.1.1959, [no collector,]. 8.

Allotype.— Figures 13B, 14B. CAS, type no. 15176. Mexico:
Baja California Sur, first wash on road to Miraflores off Mex[ico]
H[igh]w[a]y 1, 24.IV.1979, D. B. Weissman [= Stop 19] 79-95.
9.

Paratypes.— CAS; LEM, type no. KPY-21-06-P3, Mexico:
same data as allotype, 2 2 9; SDSNH, Mexico: Baja California
Sur, 5.2 mi. W of Hwy. 1, road to Coro [or Coron, just S of Los
Bariles and N of Buena Vista], 23.111.1986. D. K. Faulkner and
N. Bloomfield. 8 .

Recognition. — Small spine or sharp denticle present on the
inferoposterior angle of each lateral pronotal lobe (Figure 2C,
3B); integument only moderately, not strongly, rugose; tegminal
vestiges narrow; dorsal ovipositor valves tapered.

Etymology. — Bearing spines, referring to the spine or sharp
denticle on the inferoposterior angle of each lateral pronotal lobe.

Description of holotype. — Body moderately elongate and
rather slender; integument coarsely rugosostriate throughout, with
a few, small, scattered, pustular tubercles on head and pronotum;
hairs of frons, sternum, legs, and ends of abdomen neither long
nor dense.

Antennae comparatively long for the genus, i.e.. distinctly
longer than head and pronotum together, with 13 articles in
addition to scape and pedicel; each article, except for basal ones,
about twice as long as wide.

Head (Figures 1C, 2C) twice as long as basal width, distinctly
longer than pronotum; eyes large and prominent, oval, depth
about two-thirds of length; interocular space at narrowest about

two-thirds of greatest dorsal width of an eye; fastigium of vertex
bluntly elongate-triangular with almost straight margins, about
1 .7 times as long as basal width, with a pair of strong mediolateral
longitudinal ridges; median sulcus between dorsal areolae
extending backward for more than one-third of length of
fastigium; vertex proper rather abruptly convex between eyes;
median cannula distinct only in front of narrowest part of
interocular space, obsolescent behind this, a pair of strongly
raised ridges margining the posterior two-thirds of eyes; frontal
profile strongly oblique, concave and distinctly sinuous; fastigium
of frons slightly sinuous in outline, meeting fastigium of vertex at
a somewhat acute angle; frontal ridge very prominent above
antenna] bases, less strong below and narrowly sulcate to clypeus;
lateral frontal carinae distinct, sinuous, passing very close to
ventral margins of eyes, subparallel in anterior view.

Thorax (Figure 2C): Pronotum subcylindrical; disk with
median carina weak and interrupted in front of median transverse
sulcus, obsolescent behind it; lateral carinae obsolescent in front
of median sulcus, obsolete behind it; anterior margin subtruncate,
weakly concave in the middle; posterior margin weakly biarcuate:
posterior transverse sulcus weakly biarcuate, crossing disk at
about four-fifths of pronotal length; median transverse sulcus
almost straight, crossing disk behind its middle; lateral pronotal
lobe considerably longer than deep with almost straight margins;
anterior margin oblique; posterior margin vertical but very
slightly concave, bearing in its lower parts a pair of prominent
low elongate tubercles; inferoposterior angle almost a right angle,
bearing ventrally a sharp spine-like denticle. Mesonotum
transverse, about as long as distance between median and
posterior transverse sulci of pronotal disk. Metanotum quadrate;
metathoracic epimera very narrow, elongate, tapered anteriorly
(rather similar to, but narrower than, condition shown for female
in Figure 5B); prosternal tubercle stout, broadly conical, rounded
apically, flattened anteriorly: mesosternal lobes almost twice as
long as wide, their interspace narrow but distinct, less than half
width of a mesosternal lobe.

Wings (Figure 2C): Tegminal vestiges very small, narrowly
elongate, not extending beyond posterior margin of mesonotum;
hindwing vestiges absent or, if present, not visible beyond
posterior margin of pronotum.

Legs: Front femora barely as long as pronotum, weakly
incrassate and feebly carinate; middle femora shorter, slightly
carinate; hind femora slender, a little more than five times as long
as greatest width, not quite extending to apex of abdomen.

Abdomen: Median dorsal carina strong near posterior
margins of segments, weaker in front of these; tympana lacking;
terminalia as in Figures 4B and 5B; tergum X with posterior
emargination of moderate width, concave, dorsolateral processes
triangular; epiproct rather narrowly triangular, distinctly longer
than wide; cerci somewhat incurved, rather slender, extending as
far as apex of epiproct; subgenital plate in lateral view acute.

Phallic structures (Figures 6C, 7C): Epiphallus small, with
comparatively narrow bridge but broad lateral plates; posterior
emargination broadly and deeply concave; lophi short and
dorsally directed; lateral appendices comparatively short and
stout, their subterminal processes prominent but distinctly shorter
than rest of appendices; endophallus relatively small, with apical
parts of aedeagal valves and sclerites moderately elongate and
slender, somewhat awl-like, a little longer than basal parts of

22

D. K. McE. Kevan

valves.

Coloration more or less uniformly grayish brown but with
frons in region of median ocellus, most prominent pans of median
dorsal carinae of head, abdomen, posterior (inner) faces of middle
legs, and anterior (ventral) margins of tegminal vestiges blackish;
rest of tegminal vestiges, ventral regions, and inner faces of hind
femora pale, more or less testaceous; outer faces of hind tibiae
brownish; genae and lower pans of lateral pronotal lobes without
a pale stripe.

Measurements (mm); Length of body 18.5; antenna 7.5; head
3.3; pronotum (mid-dorsal) 2.6; hind femur 10.3; maximum width
of hind femur 1.9.

Description of Allotype. — This specimen agrees in general
with the holotype but differs in its larger size and somewhat more
robust build and in the abdominal terminalia. Other differences
are as follows:

Body with more numerous pustular tubercles.

Antennae: More ensiform. about equal in length to head and
pronotum together, with distinctly broader and relatively shorter
articles.

Head (Figures ID and 2D) a little shorter in relation to width,
about 1.7 times as long as basal width, very little longer than
pronotum; eyes less prominently and more elongately oval,
greatest depth less than two-thirds of length; interocular space, at
narrowest, considerably wider than greatest dorsal width of an
eye; fastigium of vertex relatively broader, only about 1.25 times
as long as basal width; vertex proper only weakly convex between
eyes; frontal profile slightly less oblique and scarcely sinuous;
lateral frontal carinae a little more divergent in anterior view.

Thorax: Pronotum slightly divergent posteriorly in dorsal
view, anterior margin of disk straight in middle: spine-like
denticle of inferoposterior angle of lateral pronotal lobe longer
and forming a more definite spine on left side; metanotum slightly
transverse; metathoracic epimera slightly wider (Figure 3B);
prosternal tubercle a little more transverse and less conical;
mesosternal lobes not twice as long as wide, their interspace
distinctly more than half the width of a lobe.

Wings: Tegminal vestiges extending slightly beyond posterior
margin of mesonotum. (Figure 3B is of a paratype with shorter
tegmina).

Legs: Front femora shorter than pronotum and not incrassate;
hind femora barely surpassing abdominal tergum VI.

Abdomen: Median dorsal carina strong throughout;
terminalia as in Figure 8B; epiproct a little broader than in male;
cerci short, conical, ovipositor with dorsal valves tapering in
lateral view (Figure 8B).

[Concealed copulatory structures (from LEM paratype KPY-
21-06-P3): Subgenital plate dorsally (Figure 9B) with egg-guide
acute, triangular, posterior margin on either side of it with short
longitudinal striae extending for about half width of margin;
columellae comma-shaped, not very heavily sclerotized; weakly
sclerotized patches associated with them subquadrate;
spermalheca as in Figure l()B.|

Coloration generally tawny brown (including hind tibiae) with
some mottling, notably on outer faces of hind femora; portions of
sculpture flecked with sepia; tegminal vestiges with anterior
(ventral) margins black and posterior (dorsal) margins red.

Measurements (mm): Length of body 33.0; antenna 9.8; head
5.0; pronotum (mid-dorsal) 4.6; hind femur 12.8; maximum width

of hind femur 2.2.

Paratypes. — These agree quite closely with the holotype and
allotype, but the male (SDSNH — in better condition than the
holotype. but examined too late to be so designated) has slightly
less prominent denticle and tubercles on the inferoposterior angle
of the left lateral pronotal lobe. It is also larger, with the
following measurements (mm): length of body 23.0: antenna
10.0; head 4.0; pronotum (mid-dorsal) 3.1: hind femur 10.6;
maximum width of hind femur 2.0. One female paratype (LEM)
is 29 mm and the other (CAS) 32.5 mm long. The latter is
distinctly darker and more heavily mottled with the posterior
margins of the first four abdominal terga suffused pinkish; the
inferoposterior angles of both lateral pronotal lobes bear longer
spines. The LEM paratype is colored much as the allotype, but
the anterior (inferior) margins of the tegminal vestiges
(particularly the left one) are less black; the right lateral pronotal
lobe bears a distinct spine on its inferoposterior angle, but the left
one has only a sharp denticle. In both, the tegminal vestiges are a
little shorter than in the allotype, not surpassing the posterior
margin of the mesonotum (Figure 3B).

Distribution (Figure 1 1 ). — This species has been found so far
only in the southernmost part of Baja California Sur. south of
24°N latitude. In addition to the type series, there are also two
female nymphs (about one-third grown) that clearly belong to this
species on account of their strongly striated bodies and the
distinct spines on the inferoposterior angles of the lateral pronotal
lobes. They are not regarded as paratypes. One bears the data
"2.7 mi. SE Valle Perdido. El. 1800'. 27-29. XL 1968, E. L.
Sleeper" (LEM ex CSLB): the other is labeled "2 mi. S of El
Triunfo. 17.XI.1974," D. Otte (ANSP).

Subgenus Atyphacris Kevan, Singh, and Akbar. 1964. stat. nov.

Atyphacris Kevan. Singh, and Akbar. 1964: 233-236. 240; Kevan 1477:
66 (all previous literature cited, including that for synonyms
Calamacris Rehn {partim), Icthiacris {sic. partim), Atyphoseirtus
Bruner (nomen nudum), and Atyphoscirtula Kevan. Singh, and Akbar
(nomen nudum)]; Kevan 1978: 15.

Type species. — By original monotypy. Calamacris californica
Bruner. 1906. nee (I. Bolivar) = lc[h]thiacris aptera Hebard.
1932 (replacement name).

Redescription. — Sculpture weakly striate, somewhat rugose,
granular or punctate, but tubercles feeble or lacking; fastigium of
vertex variable, sometimes long, sometimes no longer than basal
width (Figure 1E-M); vertex proper evenly or only weakly
convex dorsally (Figure 2E-M); frontal profile strongly oblique
or not. seldom strongly or sinuously concave (Figure 2E-M);
inferoposterior region of lateral pronotal lobe not rugose, angle
region lacking a tubercle, denticle, or spine; tegminal vestiges
narrow and slip-like (Figure 3C-G) or virtually if not completely
lacking (Figure 3H).

Males: Abdominal terminalia dorsally as in Figure 4C-G;
subgenital plate in lateral view acute to blunt (Figure 5C-G);
epiphallus with posterior margin deeply U-shaped or slot-like,
anterolaterally directed subterminal process of lateral appendices
not prominent (Figure 6D-N); apical pans of aedeagal valves and
sclerites rather short to very long and slender (Figure 7D-N).

Females: Metathoracic epimera expanded dorsally or not,
without tubercles (Figure 3C-H); dorsal ovipositor valves in

The Genus Ichthiacris

23

Figure 14. Ichthiacris n. spp., type specimens, lateral view. A, /. (/.) spinifera, 6 holotype; B, the same, 9 allotype; C, /. {Atyphacris) celata, 6
holotype.

24

D. K. McE. Kevan

T I. (A.) elongata

• I. (A.) cali form' ca

■ I. (A.) celata

A I. (A.) aptera

Figure 15. Known general distribution of Ichthiacris, subgenus Atyphacris, species other than /. M.) costulata. Inverted triangles, /. (A.) elongata;
circles, /. (A.) californica; squares (inset). /. {A.) celata; upright triangles (inset), /. {A.) aptera.

The Genus Ichthiacris

25

0* 109'

Figure 16. Known distribution of Ichthiacris (Atyphacris) costulata (circles).

26 D. K. McE. Kevan

Table 3. Characters distinguishing species of Ichthiacris (Atypkacris).

Character

I. {A.) elongate

I. (A.) cosntlaiah

Known distribution

Body form
Fastigium of vertex

Frontal profile

Tegminal vestiges

Males
10th abdominal tergum (dorsal)
Posterior emargination

NW Mexican mainland
(Sonora. Sinaloa)

Extremely elongate/

Much longer than wide (Fig. IE. F)/

Extremely oblique (Fig. 2E, F)/

Slip-like, very narrow (Fig. 3C)

Rather deep (Fig. 4C>

Ba|a California, southern tip
toalittleNof28°N

Elongate

Usually distinctly longer than wide
(Fig. 1G. H)

Usually rather strongly oblique
(Fig. 2G. H)

Slip-like (Fig. 3D. E)
(occasionally very minute)

Relatively narrow, not deep (Fig. 4D)

Posterior dorsolateral projections

Triangular, rather blunt
(Fig.4C)

Not prominent, pointed (Fig. 4D)

Cerci

Very elongate, extending well beyond
apex of epiproct (Figs. 4C and 5C)'

Elongate-conical, fairly robust
(Figs. 4D and 5D)

Subgenital plate (lateralview)

Epiphallus

Posterior emargination

Endophallus (lateral)

Apical in relation to basal parts
of aedeagal valves

Females

Metathoracic epimera

Subgenital plate (dorsal)

Sclerotized areas on either
side of egg-guide

Columellae

Posterior margin on either
side of egg-guide

Spermatheca and appendix (size)

Very acute (Fig. 5C/

Narrowly U-shaped (Fig. 6D)

Very much longer (longer than rest of
endophallus). very slender (Fig. 7D)

Not expanded, long and narrow
(Fig. 3C)/

Broadly crescentic (Fig. 9C)

Moderatly strong, rather distant
from posterior margin (Fig. 9C)

Convex, weakly crenulate (Fig. 9C)

Large for genus (Fig. IOC)

Moderately acute (Fig. 5D)

Slot-like (Fig. 6E-H)

Little if any longer, slender
(Fig. 7E-H)

Slightly expanded (Fig. 3D, E)

Large, subquadrate or rectangular
(Fig. 9D, E)

Strong, rather distant from
posterior margin (Fig. 9D, E)

Somewhat biarcuate. quile distinctly
crenulate (Fig. 9D, E)

Average for genus or slightly larger
(Fig. 10D, E)

"Figures IE. F. 2E, F. 3C, 4C. 5C. 6D, 7D, 8C. 9C, IOC. 16.
''Figures 1G. H, 2G. H. 3D. E. 4D, 5D, 6E-H, 7E-H, 8D. 9D, E. 10D, E. 15.
I igures II, J. 21. J, 3F. G, 4E. 5E, 6I-L. 7I-L, 8D. 9F. G. 10F. G. 16.
''Figures IK. 2K. 4F, 5F, 6M. 7M. 13C, 14C, 16.
^Figures 1L. M. 2L. M. 3H. 4G. 5G. 6N, 7N, 8E. 9H, LOH, 16.
/More obvious external feature.

The Genus Ichthicu ris

27

I. (A.) californica0

HA.) celaki'1

I (A^aptera?

Baja California, southern tip
to a little N of 25°N

Elongate

Usually little longer than wide
(Fig. II. J)

Usually less strongly oblique
(Fig. 21, J)

Slip-like (Fig. 3F.G)

Baja California, far S: known
only from Sierra de La Laguna

Not elongate for genus/

Distinctly longer than wide (Fig. IK)

Rather strongly oblique (Fig. 2K)

Virtually (cf. Fig. 3H)or
completely absent'

Baja California S of 24°N only

Not elongate for genus/

No longer than wide (Fig. 1L. M/

Not very oblique (Fig. 2L, M)/

Virtually (Fig. 3H) or completely
absent

Usually relatively wide and rather
shallow (Fig. 4E)

Usually prominent, bluntly
triangular or lobe-like (Fig. 4E|

Relatively narrow, not deep (Fig. 4F)

Not prominent, pointed (Fig. 4F)

Relatively narrow, not deep (Fig. 4G)

Not prominent, pointed (Fig. 4G)

Moderately elongate, fairly robust
(Figs. 4E and 5E)

Elongate-conical, rather slender
(Figs. 4F and 5F)

Conical, robust (Figs. 4G and 5G/

Rather blunt (Fig. 5E)

Acute (Fig. 5F)

Blunt (Fig. 5G/

U-shaped (Fig. 6I-L)

Broadly U-shaped (Fig. 6M)

Broadly U-shaped (Fig. 6N)

Much longer, slender (Fig. 7I-L)

A little longer, rather robust (Fig. 7M)

Not longer, rather robust (Fig. 7N)

Distinctly, if quite moderately, to
very strongly expanded (Fig 3F, G)/

['.' not expanded]

Slightly expanded (Fig. 3H)

Smaller, rhomboid, wider anteriorly
(Fig. 9F, G)

Strong, not very distant from
posterior margin (Fig. 9F. G)

Slightly convex, rather less
crenulate (Fig. 9F. G)

Indistinct, triangular (Fig. 9H)

Weak, not very distant from
posterior margin (Fig. 9H)

Rather straight, weakly crenulate
(Fig. 9H)

Average for genus (Fig I0F, G)

Average for genus (Fig. 10H)

28

D. K. McE. Kevan

lateral view always tapering (Figure 8C-E); subgenital plate
dorsally with posterior margin on either side of egg-guide lacking
longitudinal striae, crenulate at most and for only a short distance
along margin (Figure 9C-H); receptacula seminis as in Figure
10OH.

Distribution: — Southern Baja California and northwestern
Mexican mainland (Figures 15 and 16).

Included species. — /. (A.) elongata Kevan, Singh, and Akbar,
1964, /. (A.) costulata Bolivar, 1905, /. (A.) californica Bolivar,
1905, /. (A.) celata, n. sp. (Figures 13C and 14C), and /. (A.)
aptera Hebard, 1932. They may be distinguished from one
another as indicated in Table 3.

Ichthiacris {Atyphacris) elongata Kevan. Singh and Akbar, 1964
(Figures 1E,F; 2E. F: 3C; 4C: 5C; 6D; 7D; 8C; 9C; IOC: 15)

Ichthiacris elongata Kevan, Singh, and Akbar, 1964: 136. fig. 1 (map)
240. 247, 248. fig. 7. 249, 292, 243 (plate III), figs. A-D; Rentz, 1965:
60 [Icthiacris {sic)]; Kevan. Akbar. and Chang. 1971: 132. figs. 9 A.
B. 134: Kevan. 1477: 65; Kevan. 1978: 5, fig. 1 (map). 15. 25: Kevan.
1981: [28].

Recognition. — Form extremely elongate, including head
(Figures IE. F; 2E, F), subgenital plate (both sexes) (Figures 4C;
5C; 9C), and male cerci (Figures 4C; 5C); female cerci also long
(see Figure 8C); receptaculum seminis diagnostic, the
spermatheca and its appendix considerably larger than in other
members of the genus (compare Figure IOC with 10A. B, and
D-G). Other distinguishing features are indicated in Table 3.

Distribution (Figure 15). — Known only by a few specimens
from the northwestern Mexican states of Sinaloa and Sonora.
Sinaloa: 26 mi. N of Pericos [type locality (Kevan et al. 1964);
Rentz 1965: Kevan 1978)]; 5 and 30 mi. N of Calican (Kevan
1978). Sonora. Alamos [allotype locality (Kevan et al. 1964;
Rentz 1965; Kevan 1978)].

Ichthiacris {Atyphacris) costulata Bolivar. 1905

(Figures 1G. H; 2G, H; 3D. E; 4D; 5D; 6E-H; 7E-H; 8D: 9D. E;

10D. E; I6l

Ichthiacris costulata Bolivar, 1905: 287. 288; Kevan. Singh, and Akbar.

1964: 236. fig. 1 (map— partim), 240. 244. 245 (partim), 246. fig. 6.

247, 249, 290. 291 (plate II). fig. A-L: Kevan. 1977: 65. 644 (both

partim) |this work cites all previous references under this name and its

synonyms, except for Hebard (19231 below|; Kevan. 1978: 5, fig. 1

(map). 15, 24 (all partim); 1981: |28| (partim).
Calamacris mexicana Brunei'. 1906: 200. 201 [for other relevant

references including synonymous combinations, except for Hebard

(1923). see Kevan (I977)|; Hebard. 1923: 326 [mexicanus(sic)]; One.

1978: 29 (type catalogue).
Calamat ris palmeri Bruner. 1906: 200. 202 [for other relevant references.

see Kevan (1977)|; One. 1978: 29 (type catalogue).
Calamai ris ot ulata Bruner. 1906: 200. 202 (lor other relevant references.

see Kevan (1977)]; Otte, 1978: 29 (type catalogue).
Ichthiacris mexicana; Hebard. 1932: 268 {partim).
Ichthiacris rehni {net Bolivar); Kevan. Singh, and Akbar, 1964; 236, fig.

1 (map), 241. fig. 4, 242 only (all partim); Kevan. 1977: 44 only

(partim — by inference, citing foregoing i; Kevan. 1978: 5. tig. 1

(map), 23. 24 (all partim).
I, hthiacris spp.: Descamps, 1976: 294 (partim).

Recognition. — This species (the most widely distributed in

Baja California) is somewhat variable and can be confused with
others, but its more elongate form and the presence of small slip-
like tegminal vestiges distinguish it from /. (A.) celata and /. (A.)
aptera (the latter also has a distinctly shorter fastigium of the
vertex and less strongly oblique frontal profile). /. (A.) costulata
lacks the rugose striations and particularly the spine or denticle on
the inferoposterior angle of the lateral pronotal lobe found in /.
(/.) spinifera. I. (/.) rehni differs from /. (A.) costulata not only in
its more rugose appearance but also in its more strongly
excavated and sinuous frontal profile and more abruptly convex
vertex, particularly of the males (see Figure 2). Females of /. (/.)
rehni also have proportionately much wider tegminal vestiges
(see Figure 3). deeper, apically sickle-like dorsal ovipositor
valves (see Figure 8). and more or less straight, not convex, dorsal
margins to the metathoracic epimera (see Figure 3). /. (A.)
costulata and /. (A.) californica (the latter known only from the
southern third of Baja California Sur) are the species most likely
to be confused with each other. Their concealed copulatory
structures are quite different (compare the two species in Figures
6 and 7), but. in general, /. (A.) costulata seems to average a little
smaller in size (though this is difficult to quantify) and usually
has a slightly longer fastigium of the vertex and more oblique
frontal profile (Figures 1 and 2). Males usually have less elongate
median antennal articles, a more acute subgenital plate as seen in
lateral view (Figure 5D), less prominent dorsolateral processes on
the posterior margin of abdominal tergum X, and slightly shorter
cerci (Figure 4D). In females, the metathoracic epimera are not
less than four times as long as wide (Figure 3D, E) and not
broadly expanded as in /. (A.) californica (Figure 3F, G).

Distribution (Figure 16). — The unique female holotype of/.
costulata was described as being from "Basse Californie,"
without more precise locality. The locality erroneously ascribed
to the unique male holotype of Calamacris oculata was "(extreme
southern) Arizona." though "probably secured by G. Eisen [and
certainly] in southern Lower California" (Hebard 1932). /. (A.)
costulata is widely distributed throughout southern Baja
California, where it occurs from the extreme tip to nearly
28°30'N latitude and from sea level to well above 1200 m
elevation. It was known previously from some of the Gulf islands
and is recorded from some Pacific coast islands. More precise
localities for material studied are given below, including those for
the unique holotypes of the two other synonymous nominal
species {mexicana Bruner and palmeri Bruner). Previously
published records are indicated by the locality and references
only.

Material examined. — Baja California Norte: "Mesquital"
(presumably meaning Rancho Mezquital. though there is a
possibility that the record might refer to El Mezquital, an
abandoned American "colony" of the 1860s. about 40 km WNW
of Santa Rosalia in Baja California Sur) [recorded as /. rehni by
Kevan el al. (1964) and Kevan (1978)]: San Francisquito Bay
| large juvenile recorded as Calamacris mexicanus (sic) by Hebard
( 1923) but not noted in subsequent literature; ANSP|.

Baja California Sur: [Rancho] Los Angeles, arroyo, 0.5 km S
of km 123 S of Guerrero Negro on Mex[ico] H[igh[w[a]y 1, Stop
39. 1 1. VII. 1978. D. B. Weissman and D. C. Lighlfoot. 1 9; 14
mi. S of El Arco (Kevan 1978); 20 mi. S of El Arco [recorded as
/. rehni by Kevan et al. ( 1964) and Kevan (1978)]; Arroyo Arco
(as "Arce") 0.64 km S of km 96 N[W] of Santa Rosalia on Mex.

The Genus Ichthiacris

29

Hwy. 1, Stop 40. 11. VII. 1978. D. B. Weissman and D. C.
Lightfoot, 1 <J, 2 9 2 ; La Burrera, wash. Stop 79-209,
22.EX.1979, D. B. Weissman et al., 1 tJ; 14 km E of Mex. Hwy. 1
on road to La Burrera. elev. 304 m. Stop 79-87. 22.IV. 1979, D. B.
Weissman. 1 6, 1 juv. (small 9. probabl) this species); Cabo
Falso, 7 km W of Cabo San Lucas, 1.1.1979. P. A. Rude. 1 6;
Cabo San |as "Cape St."] Lucas [holotype of Calamacris palmed,
see Bruner (1906). Rehn and Hebard (1912). Hebard (1932).
Kevan et al. ( 1964). Kevan (1978). Otte ( 1978)]: Cabo [as
"Cape"'| San Lucas [recorded as Calamacris mexicana by Hebard
(1931)]; Cabo San Lucas. Hotel Fmisterra. 8-14. IX. 1978, J. P.
and K. E. Donahue. 3 2 9; 7.8 km E of Cabo San Lucas at km 7.8
on Mex. Hwy. 1. Stop 79-91, 23.IV.1979. D. B. Weissman. 1 6. 1
2 , 4 juvs (29.2 very small ); 2 mi. SW of Caduafio [as
"Cuduano"]. 26.VIII.1982. D. K. Faulkner and J. Brown, 1 2
(SDSNH); ca. 24 km. S of El Cien Inear Rancho El Hilario] at km
76 on Mex. Hwy. 1. 29. VII. 1977. D. B. Weissman. 1 9 ;
Comondii [recorded as /. mexicana by Hebard (1932)]:
Coronados Island, sweeping vegetation. 16. XII. 1983. N. S. Cobb.
12,4 lst-instar juvs. (99); Coronados I. and Danzante I.
[recorded as juv. Calamacris mexicanus (sic) by Hebard (1923)
and as juv. /. rehni by Kevan et al. (1964) and Kevan (1978);
seem to be present species]; 2.5 km SE of La Huerta [about half-
way between La Paz and Punta de la Ventana] (Kevan 1978); trail
from La Laguna to La Burerra, elev. 1219-1828 m. Stop 75,
22.VII.1978. D. B. Weissman and D. C. Lightfoot, 1 6. 1 2 (both
very small); 22.0 km N of Loreto, elev. 30 m. dry thorn forest.
wash, taken on low sunflower-like composite not mature enough
to be identified. 21-22.XI.1980, E. L. Sleeper, 2612/11325. I 5
(CSLB); Loreto [recorded as large 2 juvs. of Calamacris
mexicanus {sic) by Hebard (1923) and as adult 9 of/, rehni with
same data by Kevan el al. (1964) and Kevan (1978)]; Loreto.
24.X.1974, M. Descamps. 1 9 (ANSP): Isla Magdalena. Bahia
Santa Maria, ca. 1.5 km N of Punta Hughes at Smart Peak. Stop
83-95. 9. VII. 1983. D. B. Weissman and V F. Lee, 1 6*. 1 9; Isla
Magdalena, canyon 1 km NW of Puerto [Bahia] Magdalena, Stop
83-95, 8. VII. 1983, [D. B. Weissman.] D. K. Faulkner. D. C.
Lightfoot. 3 6 6,2 9 9; Isla Magdalena, path from Smart Peak to
Laguna Santa Maria. Stop 83-96. 9. VII. 1983. [D. B. Weissman.]
D. K. Faulkner, and D. C. Lightfoot. 2 juvs. (6\ 9. half grown);
San Marcos Island [juvs. recorded a /. rehni by Kevan et al.
(1964) and Kevan (1978)]; Miraflores. in palm grove,
22.VII.1971, H. G. Real and R. E. Main, 1 9; 0.8 km W of Mex.
Hwy. 1 on road to Miraflores. elev. 210 m. Stop 79-206.
26. IX. 1979. D. B. Weissman et al.. 1 9 [very small and nymph-
like; taken with /. (A.) califomica]: 5.0 km SW of Miraflores.
elev. 520 m. Cape thorn forest, 2-3.1.1981, E. L. and K. W.
Sleeper. 2321/10948, 1 6 (CSLB): 34.5 km S of Mulege. elev. 90
m. Gulf Coast desert, on Encelia farinosa ssp.'?. E. L. and K. W.
Sleeper, 2641/1 1 155, 1 9 (CSLB); 40 km S of Mulege at 0.5 km
N of km 93 on Mex. Hwy. 1, Stop 79-76. 19. IV 1979, D. B.
Weissman. 1 6 , 1 juv. ( 2 , seems to be this sp.); Patrocinio [ca. 55
km S of San Ignacio; type of Calamacris mexicana. see Bruner
(1906), Rehn and Hebard. (1912), Hebard (1923, 1931). Kevan et
al. (1964), Kevan ( 1978), Otte ( 1978)]; 44 km W of La Paz at 0.2
km S of km 44 on Mex. Hwy. 1. Stop 79-14. 31. XII. 1978. D. B.
Weissman et al., 1 2 (taken as juv. and reared; labeled "adult
30.V.1979"); as last but 0.16 km S, Stop 79-83. 21. IV 1979, D. B.
Weissman, 1 6 ; 26 mi. W of La Paz, black/white light.

II. VIII. 1976, J. Chemsak. J. Doyen, and J. Powell. I cT; 8 km W
of La Pa/ al 0.5 km N of km 8 on Mex. Hwy. 1. Stop 7l*-S4.
21. 1 VI 979. D. B. Weissman, 2 2 2,1 juv. ( 2 ); 7 mi. SW of La
Paz. at light. 6.VIII.1966, E. and J. Lindley and P. D. Head. 1 6;
23.0 km SW of La Paz. Cape thorn forest (Kevan 1978); 12 mi.
[almost due] N of La Paz on road to Pichilingue (Kevan el al.
1964, Kevan 1978) [2 9 9,1 atypical— see Kevan et al. ( 1964:
plate II. figs. K. L). but subgenital plate confirms this species];
near Pichilingue (Kevan et al. 1964; omitted by Kevan I47S);
Playa Lobos near Todos Santos, on bushes (mainly Hymenoclea)
at night; 10.XII.1979. P. A. Rude. 1 6 [with /. (A.) califomica];
Puerto Escondido [coast opposite S end of Isla del Carmen |. Stop
79-96. 25. IV. 1979. D. B. Weissman. 1 6,1 2,1 juv. ( 5 ); 0.5 km
along road to Puerto Escondido off Mex. Hwy. 1. 28. VII. 1977. D.
B. Weissman. 1 juv. (2. ?last instar); 18 km W of San Ignacio
[along Mex. Hwy. I at] km 90, 24.111.1980. J. Pinto, 1 cT;25kmS
of San Ignacio, wash at km 49 on Mex. Hwy. 1, elev. 213 m. Stop
79-73. 18.IV1979. D. B. Weissman. 4 2 2; 12 km N of San
Pedro at km 203 on Mex. Hwy. 1. 3 1. VII. 1977. D. B. Weissman.
1 6. 1 2; 10 km S of San Pedro at km 183.7 on Mex. Hwy. 1,
31.VII.1977. D. B. Weissman, 3 6 6, 1 2 (rather small, teneral), 2
juvs. (22); [Sierra] San Lazaro [now Sierra Victoria, probably
not far NW of San Jose del Cabo: young juvs. recorded as /. rehni
by Kevan el al. (1964), corrected to present species by Kevan
(1978): might be /. (A.) califomica, but probably not]; 17.5 km N
of Santa Anita turnoff at km 64 from San Jose del Cabo on Mex.
[Hwy.] 1, Stop 67, 19. VII. 1978. D. B. Weissman and D. C.
Lightfoot. 1 6 : 27.7 mi. NE of Arroyo [de] San Miguel [ca.
26°40"N. 112°30"W], 1.IV1985. N. Bloomfield and D. K.
Faulkner, 1 2 (atypically narrow fastigium of vertex) (SDSNH):

5 km N of Santa Anita at km 53 on Mex. Hwy. 1, 1. VIII. 1977. D.
B. Weissman, 1 6; Isla Santa Margarita, washes immediately SW
of Puerto Alcatraz, Stop 83-91. 7.VII.1983, D. B. Weissman. 2

6 6 (one with karyotype no. T 83-41, became adult
28. VIII. 1983), 1 9 (small, with maturation date of 17.IX.1983);
Todos Santos, garden area. Stop 76, 22. VII. 1978, D. B. Weissman
and D. C. Lightfoot. 1 6 [with /. (A.) califomica]; 11.2 km S of
Todos Santos. Stop 79-208. 27. IX. 1979. D. B. Weissman et al.. 1
6 [with /. (A.) califomica]; [El] Triunfo and 1.3 mi. NW of El
Tfiunfo [recorded as /. rehni by Kevan et al. (1964) and Kevan
(1978); former recorded as adult 9 9 but are actually last-instar
juvs.: latter are small juvs. but seem to be present species]; 7.5 km
W of El Triunfo (Kevan 1978).

In addition to the foregoing specimens. I have more recently
examined, through the courtesy of Dr. D. B. Weissman. numerous
specimens collected from various islands in the Gulf of
California. They are listed here separately to avoid repetition of
collectors' names and explanation of code numbers. Specimens
dated 1984 were all collected by D. B. Weissman and D. C.
Lightfoot; those dated 1985 indicate D. K. Faulkner as a third
member of the team. Code numbers, where given, relate to life-
history and cytological studies. A. molt to adult stage; K. date
killed after being reared from nymph. The dates following these
letters on the data labels of the specimens are (or were) entirely in
Arabic numerals with the month before the day; they are given
here with day before month, the latter in Roman numerals, to
avoid confusion. T + number, testis sample.

Isla Danzante. Slop 84-31. 16. VII. 1984. 3 68 (A8/5. T-
84/59; A9/5; A7/30, T-84/60), 6 9 9 (A8/2; A9/1; A9/6; A9/25; 2

30

D. K. McE. Kevan

uncoded): Isla del Carmen, first major canyon S of Puma Cholla.
18.VII.1984. S[top] 84-35. 1 6*. 1 9, 1 juv. (9) (all uncoded):
Isla Partida. 9. VII. 1985, Stop 85-77, 5 6 6 (A8/31, T-85/78;
A9/11. T-85/80; A9/14, T-85/81; A9/15, T-85/82; A9/25. T-
85/83), 9 9 9 (2 X A8/31; A9/27; 2 X Kll/4: 4 uncoded); Isla
Santa Cruz. 10.VI1.1985. Stop 85-78, 2 66 (T-85/45: A9/2. T-
85-76), 2 9 9 (uncoded); Isla Cerralvo. canyon on west side near
Puma el Union, 15.VII.1985, Stop 85-85, 6 66 (T-85/46; T-
85/47: A8/2. T-85/73; 3 uncoded). 5 9 9 (K8/27; Kll/4; 3
uncoded); Isla San Jose, north end. Stop 85-80. 2.VII.1985. 1 6
(unusually large; A 10/24, T-85/86), 3 9 9(A9/21; 2 uncoded);
Isla San Marcos. Stop 85-87, 17. VII. 1985. 4 66 (T-85/40; T-
85/41 [very small]; T-85/71 [very small); T-85/72), 6 9 9 (all
very small, uncoded). 1 juv. ( 9 . uncoded); Isla del Espirito Santo.
Stop 85-75, 1. VII. 1985, 1 9 (A9/10).

Remarks. — Kevan (1977) suggested that /. californica might
prove to be synonymous with /. costulata, a suggestion also put
forward by Hebard (1932). who considered the two to be
"uncomfortably close.'* Kevan (1978) formally, but erroneously,
synonymized the species under the latter name, stating that the
phallic structures of the lectotype of /. californica are like those of
/. costulata. In fact, the two species are consistently different in
this respect (compare Figures 6E-H and 7E-H with 61-L and
7I-L). not only in the greater overall size and longer aedeagal
valves and sclerites of the latter species, but also in the form of
the posterior excavation of the epiphallus, which is wide and U-
shaped in californica. narrow and slot-like in costulata. The
erroneous synonymy resulted from a lack of male specimens for
dissection when Kevan et al. (1964) prepared their revision of
Ichthiacris. Their figure 6N (p. 246) shows the posterior
emargination of the epiphallus of /. costulata as being
considerably wider than it actually is (though not as wide as in /.
californica). Re-examination of the specimen from which the
figure was prepared revealed that the bridge of the epiphallus had
been broken across the middle, and that when the two halves were
rejoined in preparation for illustration, too generous a distance
was left between them. In the resulting figure, the posterior
emargination of the bridge appeared almost U-shaped, even if
rather narrowly so.

The difference between the two species in the form of the
metathoracic epimera of the females was not previously indicated,
but the specimen illustrated by Kevan et al. (1964: 246. fig. 6E)
shows something approaching the widest condition found in /.
[A.) costulata (cf. Figure 3E herein), which is only a little
narrower than the narrowest condition found in /. (A.) californica
(cf. Figure 3F herein). In /. (A.) costulata the epimeron may be as
wide as indicated in Figure 3E herein or as narrow as in the
alleged "/. rchni" illustrated by Kevan et al. (1964: 241. fig. 4E)
and already discussed (cf. also Figure 3D herein). In /. (A.)
californica, the female's metathoracic epimeron may be nearly
(but not quite) as narrow as shown by Kevan et al. (1964: 246.
fig. 6E, see above; also Figure 3F herein), but it is usually
considerably wider and may be almost semicircular dorsally (i.e.,
D-shaped; cf. Figure 3G herein).

There now seems to be no overlap between /. (A.) costulata
and /. (A.) californica in the width of the female's metathoracic
epimeron or in the form of the posterior emargination of the
male's epiphallus, but because of previous lack of appreciation of
the differences between the species, certain records of /. (A.)

costulata given by Kevan et al. (1964) and Kevan (1978) are
inaccurate. The specimens reported as /. costulata from the
following localities belong instead to /. (A.) californica: 3.5 mi.
W of Los Barriles; 3 mi. E of La Burrera; Corral de Piedra (Sierra
de la Taste); San Jose del Cabo; Santa Rita; 15 km E of Coloni'a
[F. de la] Toba (now Villa Insurgentes): 0.5 mi. NW of El Triunfo:
2.7 mi. SE of Valle Perdido.

Ichthiacris (Atyphacris) californica Bolivar. 1905

(Figures II. J: 21. J; 3E G; 4E: 5E; 61-L; 7I-L: cf. 8D; 9F. G; 10F,

G; 15)

/i hthiacris californica Bolivar. 1905: 287. 289; Kevan, Singh, and Akbar.

1964: 240. 288. 289 (plate 1). tig. E-H: Kevan. 1977: 64 [all previous

references cited]; Kevan. 1978: 15. 24 [incorrectly indicated as

synonym of/, costulata].
hthiacris [sic] mexicana [nee (Bruner)]; Hebard 1932: 268 (partim).
Ichthiacris costulata [nee Bolivar); Kevan. Singh, and Akbar, 1964: 261.

fig. 1 (map). 245 (only— both partim): Kevan. 1977: 65 (by

implication), 644 (both partim. latter referring to Descamps. 1976.

below); Kevan. 1978: 5. fig. 1 (map), 15. 24. 25 (all partim): 1981:

[28] (partim).
Ichthiacris spp.; Descamps, 1976: 244 {partim).

Recognition. — The differences between /. (A.) californica and
/. (A.) costulata have been indicated under that species. The
present species also resembles /. (A.) aptera when the fastigium
of the vertex is particularly short, the frontal profile thus being
even less oblique than usual. The latter species is. however, less
elongate, virtually or completely without tegminal vestiges, and
has short conical cerci in the male (as in Figures 4E and 5E) and
narrower metathoracic epimera in the female (as in Figure 3H).
The most peculiar feature of /. (A.) californica is the expansion
and "flaring" of the female's metathoracic epimera (Figure 3F. G).
This character is much more pronounced in some individuals than
in others.

Distribution (Figure 15). — The male holotype and female
allotype (the only type specimens) were described as being from
"Basse Californie" without closer indication of locality. The
species occurs only in the southernmost third of Baja California
Sur from a little N of 25°N latitude to the southern tip. from near
sea level to above 1200 m elevation. It is not yet known from any
of the offshore islands. More precise localities for the specimens
studied (other than the types) are given below. Previously
published records are indicated only by locality and reference. As
already noted. /. (A.) californica has sometimes been incorrectly
recorded in the past as /. costulata. but the converse is not the
case.

Material examined. — Baja California Sur: 3.5 km W of Los
BaiTiles [recorded as /. costulata by Kevan ( 1978)]; 3 mi. E of La
Burrera [as last]; La Burrera. wash. Stop 79-209, 27. IX. 1979, D.
B. Weissman et al.. 2 6 6. 1 9,1 juv. (9, '.'last instar) [with /.
(,4.) costulata]: 1 km W of La Burrera on road to La Burrera. Stop
79-20. 2.1.1979. D. B. Weissman et al.. 1 9; Corral de Piedra,
Sierra de la Taste [recorded by Hebard ( 1964) as /. mexicana and
"El Taste," and by Kevan et al. (1964) and Kevan (1978) as /.
costulata]; trail from La Laguna to La Burrera. 1219-1828 m.
Stop 73, 2 1. VII. 1 978, D. B. Weissman and D. C. Lightfoot, 1 9
(very small but unmistakably this species); first wash on road to
Miraflores off Mex'ieo] H[igh]w[a]y. Stop 79-27. 3.1.1979. D. B.

The Genus Ichthiat ris

31

Weissman et al., 1 9; as last, but Stop 79-95. 24.IV. 1979, D. B.
Weissman, 2 juvs. (c?.2, very small. ?this species); O.S km W of
Mex. Hwy. 1 on road to Miraflores, elev. 210 m. Stop 79-206,
26.IX.1979. D. B. Weissman et al., I 3, 4 juvs. (seem to be this
species; 1 3 appearing virtually adult except for undeveloped
tegminal vestiges and phallic structures, abdominal terminalia
more or less as adult; 3 2 2) [with /. (A.) costulata]; Playa Lobos
near Todos Santos on bushes (mainly Hymenoclea), at night,
10.XII.1972. P. A. Rude. 2 9 2(1 small. I very small with rather
oblique frontal profile though metathoracic epimera widely
expanded) [with /. [A.) costulata]; Ramal de San Antonio, 5 mi. E
of [Mex.] Hwy. 1. 7 mi. S of San Antonio, 12.X. 1983. F. Andrews
and D. K. Faulkner. 5 3 6.6 2 2 (4 of latter at "black light")
(SDSNH); 2-3 mi. SW of San Bartolo. 1.X.1981, F. Andrews and
D. Faulkner. 2 2 2 (SDSNH); San Jose de Cabo [recorded as /.
costulata by Kevan et al. (1964) and Kevan (1978)]; 6 km S of
San Pedro at km 184 on Mex. Hwy. 1, Stop 79-29. 4.1.1979, D. B.
Weissman et al., 1 3 . 2 juvs. (3 , 2, very small, presumably this
species); Santa Anita, cultivated area (milpa). Stop 79-26,
3.1.1979. D. B. Weissman et al.. 1 <J. 1 2, Santa Rita (just N of
La Salada) [recorded by Kevan (1978) as /. costulata: small but
undoubtedly this species]; 9 km N of Santiago turnoff at km 94 on
Mex. Hwy. 1, Stop 79-203. 26.IX.1979, D. B. Weissman et al.. 2
3 3: Todos Santos, garden area. Stop 76, 22. VII. 1978, D. B.
Weissman and D. C. Lightfoot, 1 2 (very small but with extra-
wide metathoracic epimera) [with /. {A.) costulata]: 11.2 km S of
Todos Santos, Stop 79-208. 27.IX.1979. D. B. Weissman et al., 1
(5,12 [with /. (A.) costulata]: El Triunfo between Cabo San
Lucas and La Paz, elev. 580 m, 16.X.1974, M. Descamps. 1 3 : El
Triunfo, 3.X.1981. F. Andrews and D. K. Faulkner, 4 3 3. 4 2 2
(all rather large; 1 2 has right metathoracic epimeron more
expanded than left); 0.5 mi. NW of El Triunfo [recorded as /. (A.)
costulata by Kevan (1978)]; 2 mi. S of El Triunfo, 17.IX.1974, D.
Otte. 1 3, 1 2 [with juv. /. (/.) spinifera]; the same, but 19 [road]
miles "S" [i.e., ca. 15 km due S], 1 2 (rather large); the same, but
29 [road] miles "S" [i.e., ca. 25 km ESE]. 1 2; 2.7 mi. SE of
Valle Perdido [recorded as /. costulata by Kevan (1978)]; Villa
Insurgentes [as Colom'a (F. de la) Toba; as last].

Remarks. — The erroneous synonymy of /. (A.) californica
under /. costulata and the incorrect records for the latter have
already been discussed under that species. The curious, flared
expansion of the female's metathoracic epimera /. californica is a
unique development, the function of which is unknown.

Ichthiacris (Atyphacris) celata. new species
(Figures IK; 2K; 4F: 5F; 6M; 7M; 13C; 14C; 15)

Holotype. — Figures 12C, 13C. CAS, type no. 15175, Mexico:
Baja California Sur. Sierra [de] La Laguna, [elev.] 1770-1850 m.
28-3 1 .VIII. 1977. E. Fisher and R. Westcott. 3 .

Paratypes. — LEM, type no. KPY-22-02-P1, Mexico: same
data as holotype, but 31. VIII. 1977 and 1. IX. 1977, respectively. 2
33.

Recognition. — Though known only from males and
immatures. /. (A.) celata is distinguishable from Ichthiacris
species other than /. (A.) aptera by its comparatively short body
and virtual to complete lack of tegminal vestiges. From the latter
species it differs in its stronger sculpture, longer fastigium of the
vertex (Figure IK), and strongly oblique frontal profile (Figure

2K). Certain small specimens of/. (A.) costulata sometimes
resemble /. (A.) celata but have clearly visible, even if minute,
tegminal vestiges, and their phallic structures (notably the
epiphallus) are quite different (Figures 6 and 7). So far as is
known, males of/, celata lack the pale stripe across the genae and
inferior parts of the lateral pronotal lobes, which are rarely absent
in males of other species of the subgenus Atyphacris. I. (A.)
celata also has rather dense longish hair, particularly on the legs
and end of the abdomen.

Etymology. — Hidden or concealed; alluding to the species'
having hitherto eluded detection.

Description of holotype. — Body not very elongate for genus,
though relatively slender, coarsely rugosopunctate throughout:
hairs of frons, sternum, legs, and end of abdomen rather long and
dense.

Antennae rather long, slightly longer than head and pronotum
together, with 13 articles in addition to scape and pedicel; except
for the basal ones, the articles are more or less alternately longer
and shorter: the longer articles (including the terminal one) at
least twice as long as wide, the shorter ones a little shorter than
this.

Head (Figures IK and 2K) not quite twice as long as its basal
width, barely longer than pronotum; eyes large and prominent,
oval, greatest depth about five-sixths of length; interocular space
at narrowest about equal to greatest dorsal width of an eye;
fastigium of vertex bluntly elongate-triangular with slightly
concave margins, about 1.2 times as long as basal width, with a
pair of strong mediolateral longitudinal ridges: median sulcus
between dorsal areolae extending backward for about one-third of
length of fastigium; vertex proper rather weakly and evenly
convex; median cannula strong, extending back to pronotum; a
pair of short, strongly raised ridges margining posterior halves of
eyes; frontal profile quite strongly oblique, distinctly concave and
very slightly sinuous; fastigium of frons almost rectangular in
outline, meeting fastigium of vertex at nearly a right angle: frontal
ridge very prominent above antennal bases, less strong below and
narrowly sulcate to clypeus; lateral frontal carinae weak but
distinct, somewhat sinuous, subparallel in anterior view.

Thorax: Pronotum subcylindrical; disk with median carina
weak but distinct throughout; lateral carinae obsolescent,
subparallel; anterior margin slightly convex; posterior
emargination broad and obtuse-angular; posterior transverse
sulcus nearly straight, crossing disk at about four-fifths of
pronotal length; median transverse sulcus slightly angulate.
crossing disk at about its middle; lateral pronotal lobe
considerably longer than deep, with almost straight margins;
anterior margin oblique, posterior margin almost vertical but
slightly concave; inferoposterior angle slightly rounded and a
little less than a right angle, bearing a few weak pustular
tubercles. Mesonotum transverse, a little longer than metazona of
pronotum and about half as long as quadrate metanotum.
Metathoracic epimera narrow, elongate, tapered anteriorly (rather
similar to condition shown in Figure 3B). Prosternal tubercle
stout, broadly conical, rounded apically. flattened and emarginate
anteriorly; mesosternal lobes about twice as long as wide, their
interspace narrow but distinct, considerably less than the width of
mesosternal lobe.

Wings: Tegminal vestiges virtually lacking, represented only
by extremely minute, very narrow pads fused with basal two-

32

D. K.McE. Kevan

thirds of lateral margins of mesonotum.

Legs: Front femora about as long as pronotum, slightly
incrassate; middle femora shorter, slightly cannate; hind femora
moderately slender, not quite five times as long as greatest width,
reaching to about end of abdomen.

Abdomen with a strong medial dorsal carina; tympana
lacking; terminalia as in Figures 4F and 5F; tergum X with
posterior emargination fairly narrow, concave; dorsolateral
processes pointed; epiproct triangular, slightly longer than wide;
cerci fairly straight and slender, reaching to near apex of epiproct;
subgenital plate in lateral view acute.

Phallic structures (Figures 6M and 7M): Epiphallus with
broad bridge and broadly U-shaped posterior emargination: lophi
rather short and almost dorsally directed: lateral appendices rather
short and stout, their subterminal processes short; endophallus
with apical parts of aedeagal valves and sclerites relatively short,
somewhat blade-like, but a little distinctly longer than basal parts
of valves.

Coloration uniformly mottled grayish brown (including genae.
entire lateral pronotal lobes, and hind tibiae) except for red inner
faces of inner femora.

Measurements (mm): Length of body 20.0; antenna 7.0; head
3.1; pronotum (mid-dorsal) 2.6: hind femur 9.0 X 2.0 mm.

Paratypes. — These agree very well with the holotype. though
they are a little smaller (19.5 and 19.0 mm long). The LEM
paratype has a slightly more sinuous frontal profile (as in Figure
2K) and lacks even a trace of a tegminal vestige on the left side;
that on the right is less than half the length of the mesonotum.
The tegminal development of the CAS paratype is similar, but it
is the right side that is completely apterous.

Distribution (Figure 16). — Known only by the type series and
a few nymphs (discussed below) from the Sierra de La Laguna in
the center of the Cape region of Baja California Sur at altitudes of
between 1770 and 2000 m.

Remarks. — In addition to the type series, there are, in CAS,
three nymphs (not regarded as paratypes). They are as follows: 1
(6. last instar). same data as holotype: 2(1 (J, very small. '.'2nd
instar; 1 2. more than one third grown). Sierra de La Laguna. La
Laguna, 17 air mi. ENE of Todos Santos, elev. 6000 feet,
12-18.XII.1979. P. A. Rude collector.

The female nymph, though relatively small, indicates that the
unknown adult female of this species would conform quite
closely to the description of the male. It would, however, have
relatively shorter antennae, a slightly wider fastigium of the
vertex, and a head shorter in proportion to the pronotum. There is
not the slightest indication of any vestigial tegmen, as there is in
comparable nymphs of the other species [except /. (A) aptera].
Though this nymph is too young for one to be ceratin, it would
seem to indicate that, in the adult female, the metathoracic
epimera. though wider than in the male, do not become
appreciably expanded dorsally and are probably similar to those
of /. (A.) aptera, as in Figure 3H. The beginnings of such
expansion are quite apparent in nymphs of/. (A.) californica at a
comparable developmental stage. This nymph also shows that the
ovipositor is almost certainly of the usual, rather slender, tapered
form. Even in quite young female nymphs of /. (/.) rehni, (he
stoutness and depth of the dorsal valves begins to be apparent.

Ichthiacris {Atyphacris) aptera Hebard. 1932
(Figures IL. M; 2L. M: 3H; 4G: 5H; 6N; 7N; 8E; 9H; 10H: 15)

Calamacris californica Bruner. 1906: 200. 201: Otte. 1978: 29 (type

catalogue).
Atyphoscirtus [nomen nudum] californicus Bruner. 1908: plate 4, figs.

24, 25a.
Icthiacris [sic] aptera. nom. now [to avoid secondary homonymy with

/. californica Bolivar. 1905] Hebard. 1932: 267.
Atyphacris californica [reverting to Bruner's specific name with

elimination of secondary homonymy, now recurrent|; Kevan, Singh,

and Akbar, 1964: 235, 236. fig. 1 (map). 237, fig. 2, 238. fig. 3, 288,

289 (plate 1). figs. A-D; Kevan. 1977: 66 [citing all previous

references including synonymous combinations]; Kevan. 1978: 5. fig.

1 (map). 15. 25; Kevan. 1981: [28].

Recognition. — /. (A.) aptera is readily separated from other
species by its comparatively short body and virtually apterous
condition [shared only with /. (A.) celata], combined with a short
fastigium of the vertex, less oblique frontal profile (Figures IL.
M; 2L. M) and short conical male cerci (Figures 4G. 5G). The
egg-guide (Figure 9H) is less acutely pointed than in other species
of the subgenus Atyphacris (Figures 9C-G) [except possibly for /.
(A.) celata. of which no adult female is known].

Distribution (Figure 15). — /. (A.) aptera is positively known
only from the Cape region of Baja California Sur. south of 24°N
latitude, from at or near sea level to at least 550 m elevation.
There is. however, a questionable, more northerly record (see
below under Remarks). Localities from which specimens have
been examined are given below. Previously published records are
given by locality and reference only.

Material examined. — Baja California Sur: La Burrera, 12 air
mi. ENE of Todos Santos, elev. 1700 feet, II. XII. 1979. P. A.
Rude, 1 6.2 9 2 (1 copulating with d); 3 mi. E of La Burrera
(Kevan 1978): San Jose del Cabo [type locality] (Bruner 1906,
Rehn and Hebard 1912. Hebard 1931, 1932. Kevan et al. 1964.
Kevan 1978, Otte 1978); (Sierra) San Lazaro (now Sierra de la
Victoria), probably not far NW of previous locality (Kevan et al.
1964. Kevan 1978); ?San Pedro (Kevan et al. 1964, Kevan
1978 — see Remarks below).

Remarks. — Although /. (,4.) aptera can be completely
apterous, it usually has minute traces of tegminal vestiges fused
with the mesonotum and partially or wholly masked by the
posterior margin of the pronotum.

There is no satisfactory evidence that this species occurs
farther north than is indicated above, but there is a questionable
record from San Pedro Martir. This was taken by Kevan el al.
(1964) to be mislabeled or confused with San Pedro in southern
Baja California Sur. San Pedro Martir is the name of a mountain
range and a well-known mission in central Baja California Norte,
apparently much too far north for this species. Kevan (1978)
suggested that this locality should not be ruled out altogether, but
no member o( the Ichthiacridini (nor any other pyrgomorphid) has
been discovered as far north as 29°N. even after recent extensive
collecting throughout Baja California. The southern locality is.
therefore, tentatively accepted here, although even that may prove
to be too far north.

The Genus l< hthiacris

33

DISCUSSION

Judged by the present degree of tribal and generic diversity
(not necessarily by the richness of species) in southeastern Asia
and Madagascar, the Pyrgomorphidae probably had their origins
in eastern Gondwanaland. The Ichthiacridini are perhaps the
most primitive of the New World members of the family because
the apical parts of the aedeagal sclerites and valves are joined to
the main part of the endophallus by a constricted "flexure" (see
Figure 7). This is a plesiomorphic character, as it is found in a
wide range of Acridoidea, but has been lost in most other
Pyrgomorphidae. except for a few Oriental and Malagasy genera.
The other New World tribes do not possess this character, but,
like them, the Ichthiacridini probably reached the Americas from
the Orient by a northern route, as. it is believed, did members of
some other groups of organisms. This would have occurred, from
an educated guess, no later than the Cretaceous Period.

As noted in the Introduction, the Ichthiacridini include, in
addition to Ichthiacris, the monotypic genera Sphenacris, from
central and east-central Mexico, and Calamacris, from central and
west-central Mexico. In both, the body is more fusiform than in
Ichthiacris and is beset with fine, more regular, pustule-like
tubercles. If the Pyrgomorphidae are considered as a whole, these
characters appear to be more primitive than those of Ichthiacris.
In Sphenacris, the tubercles are dense and tegminal vestiges are
lacking; in Calamacris, the former are sparser and the latter are
present. Of the Ichthiacridini. so far as is known, Calamacris is
geographically the most central genus.

The forebears of Calamacris may well have given rise to
Sphenacris to the east, and to Ichthiacris ancestors to the
northwest. Sphenacris might be deemed to have diverged from
Calamacris by developing a more densely and regularly pustulate
integument, by losing the tegminal vestiges completely, by a
narrowing of the posterior emargination of the epiphallus [very
similar to what is found in Ichthiacris {Anphacris) costulatd], and
by some reduction in the apical parts of the aedeagus.

In Ichthiacris. the integument has not the pustulate
appearance found in Calamacris or Sphenacris but is more
striated and irregularly rugose (Ichthiacris, sensu stricto) or less
strongly sculptured, accompanied by a more cylindrical body
(subgenus Atyphacris). The vestigial tegmina, as in Calamacris.
have, however, been retained in most, though virtually or
completely lost in /. (A.) celata and /. (A.) aptera. In /. (/.) rehni
and /. (/.) parva, the tegminal vestiges, at least in females, are
broader than in other species. This might be construed as a
plesiomorphic character. Two species of Ichthiacris (subgenus
Atyphacris) have developed a peculiar, undoubtedly apomorphic
character in the female: the expansion of the metathoracie
epimera, to a moderate degree in /. (.4.) costulata, exaggeratedly
in /. {A.) californica. The function of this morphological
modification is unknown. /. (/.) spinifera is also peculiar in its
development of a small projecting spine or denticle on the
inferoposterior angle of the lateral pronotal lobe. The
significance of this. too. is unknown.

The generally U-shaped form of the posterior emargination of
the epiphallus of Calamacris is found in the majority of species of
Ichthiacris. but the emargination has become slot-like (much as in
Sphenacris) in /. {A.) costulata and broader and shallower in /. (/.)
rehni and /. (/.) spinifera. though not in the closely related /. (/.)

parva. In these last three species (Ichthiacris, sensu stricto). but
especially in /. (/.) rehni, the anterolateral^ directed subtemiinal
processes of the lateral epiphallic appendices have become
enlarged or exaggerated. In the Sonoran-Sinaloan /. (A.)
elongata, the posterior emargination of the epiphallus is narrower
than in Calamacris and most species of Ichthiacris. but it is not
slot-like. In all species of Ichthiacris, the apical parts of the
aedeagus. referred to above, are longer than in Calamacris. least
so in /. (/.) rehni and /. (/.) parva, exaggeratedly so in /. (A.)
californica and, particularly, /. (A.) elongata. I. (I.) rehni and, it
seems. /. (/.) parva. have a peculiar female reproductive
character: the ovipositor, instead of being comparatively slender
and tapered in lateral view, as in all other species of the
Ichthiacridini, is stout with the dorsal valves deep and apically
somewhat sickle-shaped. This may be apomorphic and correlated
with the type of oviposition site, thought this is not established.

It can be postulated that Ichthiacris is derived either from a
less regularly pustulate, Calamacris-Mke ancestor that invaded the
Cape region of Baja California from the Mexican mainland, or
that both genera are descendants from such an ancestor of
southeastern Asiatic origin, which came from farther to the north,
but for the existence of which there is no tangible evidence. (No
American or Asiatic fossil pyrgomorphid is known.) Whichever
is the case, Ichthiacris has speciated in southern Baja California.

Ichthiacris, sensu stricto. has retained a slightly more
elongate-fusiform body form and a greater degree of rugosity than
has the subgenus Atyphacris. while the epiphallus has become
modified in two of the three species. Each has developed its own
peculiar characters, such as the stout sickle-like dorsal ovipositor
valves of /. (/.) rehni and /. (/.) parva, the diminutive size of the
latter, and the spine or denticle on the inferoposterior angle of the
lateral pronotal lobe of /. (/.) spinifera.

The subgenus Atyphacris would seem to have diverged by
becoming less strongly sculptured and more cylindrical with a
tendency toward a shorter fastigium of the vertex and a less
oblique frontal profile, as in /. (A.) californica and. more
particularly, in /. (A.) aptera. Other modifications must also have
occurred. The latter species and /. (A.) celata appear to have
become shorter, and they have virtually (or completely) lost all
trace of wing vestiges. /. (A.) costulata. and particularly /. (A.)
californica, in the female, have developed expanded metathoracic
epimera and. in the male of the former species only, a very narrow
slot-like posterior emargination of the epiphallus. It would also
appear that this subgenus later (before or shortly after the opening
of the Gulf of California) reached the Mexican
(Sonoran-Sinaloan) mainland (rather than vice versa), where it is
represented only by the specialized, greatly elongate /. (A.)
elongata. The appearance of this species suggests that it may
have become adapted to living on grasses instead of on
xerophilous shrubs and bushes, the usual habitat for other species
of the same genus.

The form of the aedeagus seems to have undergone parallel
modifications in both subgenera. It is probable that its apical
parts beyond the "flexure," in their primitive condition, would
have been moderately long, as in /. (/.I spinifera and /. (A.)
costulata. On the one hand, however, it seems to have become
exaggeratedly long in the long-bodied /. {A.) californica. and
particularly /. (A.) elongata. and. on the other hand, shortened in
the short-bodied /. (A.) celata and /. {A.) aptera. This has

54

D K McE. Kevan

proceeded further in /. (/. ( rehni and /. (/.) parva. In Calamacris
and Sphenacris, the shortening has proceeded even further, with
an accompanying weakening of the apical parts of the aedeagal
valves in the latter genus.

In summary. I postulate that, on the basis of world knowledge
of the Pyrgomorphidae. the Ichthiacridini are descendants of a
primitive line that invaded southern North America from eastern
Asia, perhaps in Cretaceous times, though there is no supporting
fossil evidence for this. From a CalamacrisAike ancestor later
present in northern Mexico, eventual descendants spread to Baja
California, resulting in the evolution of early Ichthiacris-Wks
forms. These apparently gave rise to two lineages: Ichthiacris,
sensu stricto. and the subgenus Atyphacris. The latter reached the
southernmost part of the peninsula, where it has speciated more
than has the nominotypical subgenus (which did not penetrate
quite so far south). Descendants of one of the southern forms of
Atyphacris must, in relatively recent times, it would seem, have
made its way back to the northwest Mexican mainland, where it is
today represented by /. (/A. I elongata, the only member of the
genus known to occur outside southern Baja California. On the
long-isolated Cedros Island in the northwesternmost part of the
range of the genus, it appears that an /. (/.) rehni ancestor gave
rise to /. (I.) parva.

ACKNOWLEDGMENTS

I thank Dr. D. B. Weissman and Dr. P. H. Arnaud. Jr., of the
California Academy of Sciences, San Francisco, for their
assistance in arranging for the loan of the bulk of the material
upon which this paper is based. I am also indebted to Dr. D. A.
Nickle of the United States Department of Agriculture Systematic
Entomology Section, U.S. National Museum of Natural History.
Washington. D.C.. and to Dr. D. Otte of the Academy of Natural
Sciences of Philadelphia, for the loan of certain other material,
mostly for re-examination. The line illustrations were inked for
publication by D. J. Klimaszewski, formerly of the Lyman
Entomological Museum and Research Laboratory: photographs
are by Mr. F. Genier. formerly of the same institution, and by Mr.
P. Langlois. Department of Entomology. McGill University. This
work was supported by a grant to the author from the National
Sciences and Engineering Research Council of Canada (Grant
No. A 1378).

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