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Revision of the Alutacea Group of Genus Schistocerca (Orthoptera: Acrididae: Cyrtacanthacridinae)

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The North American Alutacea Group of the genus Schistocerca is revised based on phylogenetic analysis and morphological comparison. Phylogenetic analysis suggests that Schistocerca alutacea sensu Dirsh is paraphyletic because S. alutacea albolineata sensu Dirsh is sister to S. obscura. Here, I recognize the monophyletic Alutacea Group consisting of six species: S. alutacea, S. rubiginosa, S. lineata, S. shoshone, S. albolineata, and S. obscura. Previous taxonomic confusion on these species is discussed, and their taxonomic ranks are accordingly raised from subspecies to species level. Neotypes of S. alutacea and S. rubiginosa are designated, and S. insignis is synonymized under S. albolineata syn. nov. Also, I argue for the validity of the name S. lineata and discuss the previous misapplication of nomenclature. Discussion of color variation and an identification key for all North American Schistocerca species are presented.
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SYSTEMATICS
Revision of the Alutacea Group of Genus Schistocerca
(Orthoptera: Acrididae: Cyrtacanthacridinae)
HOJUN SONG
1
Department of Entomology, The Ohio State University, Columbus, OH 43212
Ann. Entomol. Soc. Am. 97(3): 420Ð436 (2004)
ABSTRACT The North American Alutacea Group of the genus Schistocerca is revised based on
phylogenetic analysis and morphological comparison. Phylogenetic analysis suggests that Schistocerca
alutacea sensu Dirsh is paraphyletic because S. alutacea albolineata sensu Dirsh is sister to S. obscura.
Here, I recognize the monophyletic Alutacea Group consisting of six species: S. alutacea,S. rubiginosa,
S. lineata,S. shoshone,S. albolineata, and S. obscura. Previous taxonomic confusion on these species is
discussed, and their taxonomic ranks are accordingly raised from subspecies to species level. Neotypes
of S. alutacea and S. rubiginosa are designated, and S. insignis is synonymized under S. albolineata syn.
nov. Also, I argue for the validity of the name S. lineata and discuss the previous misapplication of
nomenclature. Discussion of color variation and an identiÞcation key for all North American Schis-
tocerca species are presented.
KEY WORDS North American Schistocerca, Alutacea Group, systematics, phylogeny, genitalia
THE LOCUST GENUS Schistocerca Stål is a biologically
fascinating group. The genus is famous for its African
species, S. gregaria (Forskål), perhaps one of the most
notorious insects in the world. Schistocerca gregaria,
the desert locust, is capable of forming an enormous
swarm consisting of billions of individuals through
density-dependent phase changes (Uvarov 1977,
Pener and Yerushalmi 1998). Although the well-
known desert locust is important and interesting in
many aspects, Schistocerca is in fact a New World
genus, containing about 50 species (Dirsh 1974). In-
deed, the desert locust is the only Old World repre-
sentative of the genus, and this transatlantic disjunc-
tion sparked heated debates concerning its origin
among entomologists (Kevan 1989, Ritchie and Pedg-
ley 1989). New World Schistocerca species are highly
diverse and have adapted to different environments.
For example, some species form a large swarm com-
parable with that of the desert locust (Harvey 1981).
Most Schistocerca species are, however, sedentary,
meaning that they do not form a swarm. They never-
theless have an extremely strong ßight capacity. Cer-
tain species show density-dependent aposematism,
which became a prime example of novel insight into
the evolution of warning coloration (Sword 1999).
These sedentary species also have high host prefer-
ence within nymphal stage, which is very curious
given that most grasshopper species are generalists.
Schistocerca ceratiola Hubbell and Walker, endemic to
central Florida, is one of few monophagous and noc-
turnal species of grasshoppers (Hubbell and Walker
1928).
Despite these attractive aspects of the genus, tax-
onomy of Schistocerca is surprisingly poorly under-
stood. This dichotomy can certainly be attributed to
the variability within a species and the similarity across
the species. Some widespread species, such as S. lin-
eata Scudder and S. nitens (Thunberg), have many
local populations adapted to speciÞc habitats, which
introduces difÞculty in taxonomic studies. Other spe-
cies, such as species within the Americana and Aluta-
cea Groups, are so similar externally that a careless
synonymy is quite possible. These factors coupled
with an inadequate taxonomic method are the exact
reasons why the most recent revision of the genus
(Dirsh 1974) is considered to be problematic.
In his revision, Dirsh (1974) used morphometric
characters exclusively to deÞne species, resulting in
numerous synonymies. He modiÞed the morphomet-
ric technique, initially used to differentiate the phase
of a locust species (Uvarov 1966), to characterize each
species using the proportional value of several mea-
surements of characters including length of body,
length of pronotum, length of tegmina (forewing),
length and width of hind femur, and width of head. He
created a pentagram using the proportional values and
visually compared the shape for each species. Al-
though these characters themselves are useful in dif-
ferentiating species in some cases, the method failed
to differentiate closely related species that are only
distinguishable by internal male genitalia. As a result,
Dirsh (1974) recognized 22 species and 21 subspecies.
The most problematic species is Schistocerca nitens,
1
E-mail: song.131@osu.edu.
0013-8746/04/0420Ð0436$04.00/0 2004 Entomological Society of America
under which Dirsh synonymized 16 names. No one has
yet re-evaluated his treatment. Other taxonomically
problematic species are S. americana (Drury) and S.
alutacea (Harris). Dirsh (1974) changed the rank of
any species that shared a similar pentagram, creating
11 subspecies under S. americana and 7 under S. aluta-
cea. Several workers doubted his taxonomic concept
and initiated a series of hybridization experiments to
test the biological meaningfulness of the revision
(Harvey 1979, 1981, 1982, Jago et al. 1979, 1982). These
works focused on the subspecies of S. americana, in-
cluding some of the most destructive locust species in
the world, and resulted in a reclassiÞcation that
yielded Þve species and four subspecies (Harvey
1981). However, no similar attempt has been made for
the sedentary species in North America, S. alutacea.
In this study, I re-evaluate DirshÕs synonymy based
on the morphological characters and known ecology.
Several ecological studies on North American Schis-
tocerca have been published since DirshÕs revision
(Sword and Chapman 1994, Chapman et al. 1995,
Howard 1995, Sword and Dopman 1999, Capinera et
al. 2001), and most of them use the names before the
synonymy. Here, I attempt to resolve long-standing
taxonomic confusions marred by disagreements
among orthopterists. I also test previous phylogenetic
relationships among the North American Schistocerca
species suggested by Hubbell (1960) and Dirsh (1974)
in light of the morphological phylogeny.
Color Variation in Schistocerca
Schistocerca is known as a taxonomically difÞcult
group mainly because of its color variation (Hebard
1923, Hubbell 1960). DirshÕs (1974) revision is a good
example of how color variation can affect the taxo-
nomic concept. His justiÞcation for synonymy, in ad-
dition to the morphometric technique, stemmed from
his erroneous claims to have found the intermediate
forms of all possible combinations with overlapping
geographic distributions (Dirsh 1974). Variability of
color in grasshoppers is well known and sometimes
ineffective in differentiating species (Hubbell 1960).
This is especially true for the North American Schis-
tocerca species. Hubbell (1960) analyzed the variabil-
ity of color patterns in S. lineata, S. alutacea, and S.
rubiginosa (Harris), and here I brießy review his work
and other studies (King and Slifer 1955, Chapman et
al. 1995, Sword and Dopman 1999).
The dorsal stripe, which extends from the vertex to
the anterior tip of pronotum, is one of several variable
color patterns in Schistocerca (Rehn 1901, 1902). In
early 1900, S. alutacea was known to be always striped,
whereas S. rubiginosa was thought to be unstriped.
Rehn (1901, 1902) observed a striped form and an
unstriped form in copula and suspected that S. rubigi-
nosa was just an unstriped form of S. alutacea. After
examining a large number of specimens of both spe-
cies identiÞed by unique phallic morphologies, Hub-
bell (1960) concluded that S. alutacea is always
striped, and S. rubiginosa can be either striped or
unstriped. He then plotted the occurrence of a dorsal
stripe in S. rubiginosa against the collecting localities
of the specimens and found that the southern popu-
lations gradually lose the dorsal stripe. Similar phe-
nomenon is found in S. shoshone (Thomas). Originally,
the species was described based on the striped form,
and Scudder (1899) described two additional species
based the absence of the dorsal stripe. Hebard (1935),
when synonymizing unstriped S. venusta Scudder un-
der striped S. shoshone, hypothesized that S. venusta
was nothing more than an unstriped form of S. shos-
hone. Chapman et al. (1995) suggested that these two
forms are associated with different host plants. Indi-
viduals that feed on jojoba, Simmondsia chinensis
(Link) C. Schneid (Buxaceae), were always striped,
and individuals that feed on mesquite, Prosopis velu-
tina Wooton (Caesalpiniaceae), were always un-
striped. Chapman and Sword (unpublished data) per-
formed several crossing experiments between striped
and unstriped forms and found that offspring were
always striped regardless of the sex of the parents,
conÞrming HebardÕs (1935) hypothesis on the vari-
ability of the stripe.
Hubbell (1960) also studied regional variations in
body color, markings of head and thorax, mesepimeral
stripe, tegminal maculation, and hind tibiae. He dem-
onstrated that different populations of a single species
could vary considerably in color patterns and that
color alone could not be used as a deÞnite diagnostic
character. For example, he showed that color of the
hind tibiae varied according to the populations, espe-
cially in S. lineata. The hind tibiae were yellowish or
brownish in the eastern populations, coral pink or red
in the midwestern populations, and black in the south-
western populations. A similar phenomenon can be
observed in S. albolineata (Thomas), whose Arizona
populations have red hind tibiae, whereas Texas and
Mexican populations have black hind tibiae (unpub-
lished data). King and Slifer (1955) found that in
Melanoplus mexicanus mexicanus (Saussure), color of
the hind tibiae was determined by the Mendelian
genetics of an autosomal gene, which could explain
the color variation of hind tibiae in Schistocerca.
Schistocerca lineata is one of the most polymorphic
species in the genus, and Hubbell (1960) showed that
coloration intensiÞes toward southwestern regions.
Populations in Illinois, Indiana, Michigan, and Ohio
were lightly colored, whereas those in Oklahoma and
Texas were conspicuously colored. He hypothesized
that the variation could be associated with food pref-
erence of a local population and was not necessarily
related to the habitat. Dirsh (1974) proposed that the
conspicuously colored form was a hybrid between S.
albolineata and S. lineata, which he used as a reason for
his synonymy. However, this Texas population is not
a hybrid, but an aposematic population of S. lineata
that probably feeds on toxic Ptelea trifoliata L. (Ru-
taceae) (Sword and Dopman 1999). Sword and Dop-
man (1999) studied several populations of S. lineata in
Texas that used different host plants as a nymphal
instar. They showed that the intensity of density-
dependent aposematism was different in populations
associated with toxic P. trifoliata and populations as-
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 421
sociated with palatable Rubus trivialis Michaux (Ro-
saceae). When crowded, nymphs from both popula-
tions produced an aposematic coloration, but Ptelea-
associated nymphs were more intense in color. Sword
(2002) suggested that the nymphal coloration associ-
ated with a host plant is genetic, not necessarily re-
lated to host plant, because he was able to rear dif-
ferent color phenotypes on the same standard diet
(lettuce). The ontogenetic host specialization in re-
lation to the aposematism is only known in Texas
populations of S. lineata, and given the speciesÕ wide
distribution, midwestern and eastern populations
might have different ecologies (Sword and Dopman
1999).
North American Schistocerca
There are 10 Schistocerca species in North America
north of Mexico (Table 1). Recently, Stidham and
Stidham (2002) reported the occurrence of S. cam-
erata Scudder in southern Texas, but because it is
mainly distributed in Mexico, it is not treated in this
revision. Hubbell (1960) recognized six species groups
in North America: Americana, Nitens, DamniÞca,
Ceratiola, Obscura, Shoshone, and Alutacea Groups.
However, this grouping is not based on a modern
cladistic analysis, and each group might not represent
a monophyletic clade. Here, I present a slight modi-
Þcation of HubbellÕs grouping and tentatively recog-
nize Þve species groups (Table 1).
The Americana Group contains large grasshopper
species including all three swarming species: the Cen-
tral American locust S. piceifrons (Walker), the South
American locust S. cancellata (Serville), and the desert
locust S. gregaria. S. americana is the only North Amer-
ican representative, distributed from Florida to Texas.
It occasionally forms minor swarms that cause damage
in orange orchards in Florida (Harvey 1981, Capinera
et al. 2001). The Nitens Group is a taxonomically
problematic group, and perhaps contains more than 10
species. S. nitens nitens sensu Dirsh (1974) is extremely
widespread and spans from the western United States
to Brazil. I examined a large number of specimens
bearing the identiÞcation label by Dirsh as S. nitens
nitens and was able to distinguish several distinct forms
based on color, male cerci, and male phallic complex.
It is not clear why he synonymized so many species
under S. nitens nitens, most of which were soundly
recognized by earlier taxonomists (Hebard 1923, 1924,
1932). North American S. nitens can be characterized
by tectiform pronotum and mottled tegmina. It is
sometimes called by its old name S. vaga (Scudder). Its
distribution ranges from southern California to Mex-
ico, with a population found in Hawaii (Bianchi and
Kajiwara 1966, Sugerman 1966). The Ceratiola Group
contains single species, S. ceratiola, which is endemic
to Central Florida. It is unique in nocturnal habit as
well as monophagy on Florida rosemary, Ceratiola
ericoides Michaux (Hubbell and Walker 1928, Franz
and Franz 1989). The DamniÞca Group also contains
one species, S. damnifica (Saussure), distributed in the
eastern United States. This species is highly homoge-
neous in form and can be easily identiÞed by having
rusty brown color, highly tectiform pronotum, and
slightly ensiform antennae. In the past, two subspecies
of S. damnifica were described: the nominal subspe-
cies and S. damnifica calidor (Rehn and Hebard). The
only difference between the two is size, and here I
follow DirshÕs (1965) synonymy in recognizing a sin-
gle species with size variability. The remaining six
species forming the Alutacea Group are subject of this
study.
Materials and Methods
In this revision, I examined six species forming the
Alutacea Group. These are S. alutacea (Harris), S.
rubiginosa (Harris), S. lineata Scudder, S. shoshone
(Thomas), S. albolineata (Thomas), and S. obscura
(Fabricius). I directly compared all known type spec-
imens with the original descriptions and DirshÕs revi-
sion.
This work was based on the study of specimens (n
7,882) from the following institutions: Academy of
Natural Sciences, Philadelphia, PA (ANSP); Brigham
Young University, Arthropod Collection, Provo, UT
(BYU); Reginald ChapmanÕs personal collection
(CHAP); Colorado State University Insect Collection,
Fort Collins, CO (CSUC); Illinois Natural History
Survey, Champaign, IL (INHS); University of Wis-
consin Insect Research Collection, Madison, WI
(IRCW); McGill University, Lyman Entomological
Table 1. Taxonomic concept changes in North American Schistocerca species
Species
Group
Currently recognized
species
Original
Description
Scudder
(1899)
Hubbell
(1960)
Dirsh
(1974)
Harvey
(1981)
Americana S. americana (Drury, 1773) Gryllus americanus S. americana S. americana S. americana americana S. americana
Nitens S. nitens (Thunberg, 1815) Gryllus nitens S. vaga S. vaga S. nitens nitens N/A
Ceratiola S. ceratiola (Hubbell and
Walker, 1928)
Schistocerca ceratiola N/A S. ceratiola S. ceratiola N/A
DamniÞca S. damnifica (Saussure, 1861) Acridium damnificum S. damnifica S. damnifica S. damnifica N/A
Alutacea S. alutacea (Harris, 1841) Acrydium alutaceum S. alutacea S. alutacea S. alutacea alutacea N/A
S. rubiginosa (Harris, 1862) Aciridium rubiginosum S. rubiginosa S. rubiginosa S. alutacea rubiginosa N/A
S. lineata Scudder, 1899 Schistocerca lineata S. lineata S. lineata S. alutacea lineata N/A
S. shoshone (Thomas, 1873) Acridium shoshone S. shoshone S. shoshone S. alutacea shoshone N/A
S. albolineata (Thomas, 1875) Acridium albolineata S. albolineata S. albolineata S. alutacea albolineata N/A
S. obscura (Fabricius, 1798) Gryllus obscurus S. obscura S. obscura S. obscura N/A
422 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
Museum, Ste. Anne de Bellevue, QC, Canada
(LEMQ); New Mexico State University Arthropod
Museum, Las Cruces, NM (NMSU); Oklahoma State
University, K.C. Emerson Museum, Stillwater, OK
(OSEC); Ohio State University Insect Collection, Co-
lumbus, OH (OSUC); University of Kansas, Snow En-
tomology Collection, Lawrence, KS (SEMC); Univer-
sity of Arkansas Arthropod Museum, Fayetteville, AR
(UARM); University of Michigan Museum of Zoology,
Ann Arbor, MI (UMMZ); University of Minnesota
Insect Collection, St. Paul, MN (UMSP); University of
Nebraska State Museum, Lincoln, NE (UNSM); Uni-
versity of Idaho WFBarr Entomological Museum,
Moscow, ID (WFRM). HubbellÕs genitalia collection
from UMMZ was also studied. All existing type spec-
imens were examined from the following institutions:
Academy of Natural Sciences, Philadelphia, PA
(ANSP); British Museum of Natural History, London,
U.K. (BMNH); California Academy of Sciences, San
Francisco, CA (CAS); Harvard University Museum of
Comparative Zoology, Cambridge, MA (MCZ); Uni-
versity of Michigan Museum of Zoology, Ann Arbor,
MI (UMMZ).
Male genitalia were extruded by inserting a probe
under the epiproct using the technique described by
Hubbell (1932). Ovipositors were dissected by making
a slit at the distal part of abdomen (Cohn and Cantrall
1974). Genitalia were placed in 10% KOH solution for
several hours to dissolve muscles. Cleared genitalia
were placed in a vial Þlled with glycerin, and each
genitalic specimen was given an identiÞcation number
to associate with the pinned specimens. All illustra-
tions were made using a drawing tube on a Wild
dissecting microscope.
To test the previously suggested phylogenetic re-
lationships, a cladistic analysis was performed using 10
North American Schistocerca species and 4 cyrtacan-
thacridine outgroup species. Acanthacris, Cyrtacan-
thacris, Ornithacris, and Halmenus (a genus endemic
to the Galapagos) were used as outgroup taxa, and
Ornithacris was used to root the tree. Twenty-two
morphological characters with a total of 55 character
states were used in the cladistic analysis (Appendix
A). Inapplicable characters were coded as Ñ.All
characters were equally weighted and treated as non-
additive. Characters 0Ð1, 6Ð8, 9Ð10, 17Ð18, 19Ð20, and
21Ð22 were coded using a mixed-coding technique.
The data matrix (Appendix B) was created in Win-
Clada (Nixon 2002), and the cladistic analysis was
carried out using NONA (Goloboff 1995). Search pa-
rameters include rs 0; hold 10,000; mult*100; and max*.
From the obtained trees, any unsupported node was
collapsed.
An electronic supplement of this study is now avail-
able (http://www.schistocerca.org/alutacea.htm).
This website includes information on each species
along with the images of all existing type specimens.
It also contains an interactive identiÞcation key, a
diagnosis, a distribution map, and a list of selected
literature. PDF Þles of the original description are also
linked to the website.
Results and Discussion
Phylogeny of North American Schistocerca
The cladistic analysis resulted in two most parsimo-
nious trees of 39 steps (Figs. 1 and 2). The strict
consensus of two most parsimonious trees collapsed
one node and produced a consensus tree of 40 steps
(consistency index [Ci] 80; retention index [Ri]
87; Fig. 3). Monophyly of the Alutacea Group was
strongly supported in by four synapomorphies, al-
though the relationship within the clade is largely
unresolved. Among the most parsimonious trees, one
topology (Fig. 1) suggests that S. alutacea and S. ru-
biginosa are sisters to S. albolineata S. obscura clade
by having constricted basal eminence of cingulum.
The alternative topology (Fig. 2) suggests that they
are sisters to S. lineata S. shoshone clade supported
by a pair of tubercles on male epiproct. Current data
alone are insufÞcient in resolving this problem, but it
is clear that the species in the Alutacea Group are very
closely related.
Figs. 13. Phylogeny of the North American Schistocerca species. (1 and 2) Two equally parsimonious cladograms
produced from the analysis (L 39; Ci 82; Ri 89). (3) Strict consensus cladogram (L 40; Ci 80; Ri 87). Four
cyrtacanthacridine outgroup taxa were collapsed as a single terminal in (1 and 2). Numbers above node are Bremer support
values, and numbers below are the number of synapomorphies for that node.
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 423
Present analysis (Fig. 3) contradicts the previously
suggested phylogenies by Hubbell (1960) and Dirsh
(1974). Hubbell (1960) suggested that S. alutacea, S.
rubiginosa, and S. lineata are sibling species, distin-
guishable by only the subtle differences in genitalic
morphologies. The current phylogeny suggests that S.
lineata is in fact sister to S. shoshone and are united by
inßated fore and middle femora. Dirsh (1974) did not
synonymize S. obscura under S. alutacea because he
considered the morphological differences to be sufÞ-
cient enough. However, the present phylogeny sug-
gests that S. obscura is sister to S. albolineata, which
was considered as a subspecies of S. alutacea by Dirsh
(1974). This result indicates that S. alutacea sensu
Dirsh is paraphyletic, thus demonstrating the inade-
quateness of his synonymy.
North American Schistocerca species are ecologi-
cally sedentary, perhaps with an exception to S. ameri-
cana. Although they possess a strong ßight capacity,
they mostly stay in the same habitat in their lifetime.
Mass migration observed in other swarming species
has never been reported in the species of the Alutacea
Group. Occasionally, aggregation behavior has been
observed in some species, causing minor damage to
crops, but they do not have the characteristics of true
swarming behavior such as cohesive ßight or nymphal
band movement (Chapman et al. 1995). Interestingly,
however, the sedentary species of the Alutacea Group
was recently found to possess a character, density-
dependent polyphenism, known to be crucial in the
swarming Schistocerca species (Sword 1999, 2002).
The desert locust S. gregaria is usually solitary, but
when population density increases, it behaves gregar-
iously followed by changes in color patterns and
shapes (Uvarov 1966). Although the sedentary species
do not behave gregariously or change in morphology,
they do change color in response to high density
(Sword 1999). Sword (1999) discovered that the
nymphs of S. lineata in Texas change color when
reared in crowded condition. Nymphs associated with
the toxic Ptelea were green when reared in isolation
but turned bright yellow and black when reared in
crowdedness. Sword (2002) later suggested that the
density-dependent polyphenism in S. lineata is par-
tially genetic, not necessarily related to the host plant
the individuals feed on. Although not studied, it is
possible that the closely related species in the Aluta-
cea Group might have the similar polyphenism. This
would indicate then that the density-dependent
polyphenism is a phylogenetically informative char-
acter.
Another interesting point deduced from the phy-
logeny is the evolution of feeding habit. Most grass-
hopper species are polyphagous (Chapman 1990).
Sword and Dopman (1999) discovered that the Texas
populations of S. lineata were monophagous as
nymphs but polyphagous as adults. The host associa-
tion is also found in the Arizona populations of S.
shoshone (Sword and Chapman 1994), and in the Texas
populations of S. obscura (Chapman and Sword, un-
published data). The phylogeny suggests that S. cera-
tiola is basal to all the other sedentary North American
species. S. ceratiola is one of few monophagous grass-
hopper species that specializes on C. ericoides (Hub-
bell and Walker 1928, Franz and Franz 1989). There-
fore, it is possible that the common ancestor between
S. ceratiola and other sedentary species evolved the
intimate hostÐplant association. The extreme
monophagy found in S. ceratiola is perhaps the result
of the intensiÞed dependency on the host plants. Be-
cause of this dependency, S. ceratiola might have de-
veloped the nocturnal habit to avoid predators. The
ancestor of the Alutacea Group could have explored
other niches but nevertheless retained the innate ten-
dency to associate with the host plants in the local
population. The fact that different local populations of
S. lineata and S. shoshone have different host associa-
tions indicates that the nymphal monophagy is rather
plastic in terms of choosing the host plants.
Table 2. Diagnostic characters, distribution, phenology, preferred habitat, and known host plants for the species in the Alutacea Group
Species Diagnostic characters Distribution Phenology Preferred habitat Known host plant
S. alutacea Highly convex rami of cingulum;
always with a yellow dorsal
stripe
Eastern United States MayÐAug Shrubby, moist to wet
(Northeastern
U.S.); dry, sand hill
(Florida)
Vaccinium
macrocarpon
S. rubiginosa Slightly inßated head; cylindrical
pronotum; small male cerci
Southeastern United
States
MayÐAug Xeric, sandy soil Unknown
S. lineata Basal eminence broad in the
middle; inßated male fore and
middle femora
Entire North America MayÐOct Variable Ptelea trifoliata, Rubus
trivialis, Quercus sp.,
Glycyrrhiza lepidota,
Taraxacum sp.
S. shoshone No marking on abdominal tergites Western U.S. JuneÐAug Riparian, desert Simmondsia chinensis,
Prosopis velutina
S. albolineata Male cerci with lower apical
angle protruding more than
upper
Southwestern United
States and Mexico
Aug-Oct Dry mountain Cercidium
microphyllum,
Gossypium hirsutum
S. obscura Male subgenital plate highly
ßared outward
Eastern United States JulyÐSept Open woodlands Celtis laevigata
424 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
The Alutacea Group
The Alutacea Group consists of six closely related
species: S. alutacea (Harris), S. rubiginosa (Harris), S.
lineata Scudder, S. shoshone (Thomas), S. albolineata
(Thomas), and S. obscura (Fabricius) (Table 2). It is
characterized by four synapomorphies: round apex of
male subgenital plate, rectangular male cerci, pres-
ence of small granules on pronotum and thorax, and
constricted basal eminence of zygoma in male phallic
complex. Useful plesiomorphic characters include un-
usually long antennae especially in males, robust body
form, tegmina just slightly longer than the tip of ab-
domen, and tegmina colored in olive green or brown.
The group is geographically conÞned in North Amer-
ica. Ecologically, the species in the group are arboreal
and found in various woody habitats (Squitier and
Capinera 2002a, Sword and Chapman 1994).
The Alutacea Group in the present sense is broader
than in the sense of Hubbell (1960). He originally
included three species, S. alutacea, S. rubiginosa, and
S. lineata, within his Alutacea Group that were united
by somewhat superÞcial characters. In this study, I
have retained the species group name, but it now
includes HubbellÕs (1960) Alutacea, Obscura, and
Shoshone Groups. Subspecies of S. alutacea sensu
Dirsh (1974) are now considered valid species, with
the exception of S. insignis Hebard, which is now a
synonym of S. albolineata.
Key to North American Schistocerca Species
1. Mesosternum with lateral lobes longer than
wide. Prosternal process always present. Male
subgenital plate always bilobed. Usually mod-
erate to large size ......2,Schistocerca Stål
Mesosternum with lateral lobes as wide as long or
wider than long. Prosternal process present or
absent. Male subgenital plate variable. Usually
small to medium size. .....Other Acrididae
2. Small for the genus (TL 29Ð32 mm; TL
39Ð41 mm), eyes highly prominent. Tegmina
narrow, antennae much longer than the length
of head and pronotum. Hind femur with two
dark dorsal bands. Taken alive, color green.
Museum specimens brown. Endemic in Cen-
tral Florida, only found on Florida rosemary
Ceratiola ericoides..................
..........S. ceratiola Hubbell and Walker
body size moderate to large. Tegmina broad .
............................. 3
3. Tegmina much longer than the tip of abdomen,
and posterior margin of tegmina colorless and
transparent. Entire tegmina with distinct black
patches or mottling ................4
Tegmina slightly longer than the tip of abdomen,
and the entire tegmina brown, olive green,
liver brown, or rusty brown, with or without
slight mottling ....................5
4. Tegmina transparent with large distinct dark
patches. Overall brown in color. Lateral lobe
of pronotum with two dark markings. Median
carina of pronotum low and prozona slightly
constricted. Antennae as long as the length of
head and pronotum. Hind femur without dor-
sal bands. Males slightly smaller than females.
................ S. americana (Drury)
Tegmina transparent with many irregular mot-
tling. Overall gray in color. Lateral lobe of
pronotum, with single dark marking and white
marking at the base. Median carina of prono-
tum slightly tectiform, especially in females
and prozona not constricted. Antennae
slightly longer than the length of head and
pronotum. Hind femur with two dorsal bands.
Males much smaller than females ........
................. S. nitens (Thunberg)
5. Entire body rusty brown sometimes with slight
mottling on tegmina. Median carina of prono-
tum distinctly elevated. Antennae shorter than
the length of head and pronotum, and weakly
ensiform .........S. damnifica (Saussure)
Body color variable with tegmina uniformly
brown or with slight mottling. Median carina
Figs. 427. External morphological characters of the
species in the Alutacea Group. (4Ð7) S. alutacea; (8Ð11) S.
rubiginosa; (12Ð15) S. lineata; (16Ð19) S. shoshone; (20Ð23) S.
albolineata; (24Ð27) S. obscura; (4, 8, 12, 16, 20, and 24) male
epiproct; (5, 9, 13, 17, 21, and 25) male subgenital plate; (6,
10, 14, 18, 22, and 26) male cercus; (7, 11, 15, 19, 23, and 27)
hind femur pattern. Bar a 1 mm for 4, 8, 12, 16, 20, and 24.
Bar b 1 mm for 5, 9, 13, 17, 21, and 25. Bar c 1mmfor
7, 11, 15, 19, 23, and 27.
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 425
of pronotum not elevated. Antennae, espe-
cially in males, much longer than the length of
head and pronotum, and Þlliform .......
................ 6,theAlutacea Group
6. Male epiproct without a pair of tubercles (Figs.
20 and 24). Apical lobes of male subgenital
plate slightly or much ßared outward (Figs. 21
and 25). Integument highly setose .......7
Male epiproct with a pair of tubercles (Figs. 4, 8,
12, and 16). Apical lobes of male subgenital
plate not ßared outward (Figs. 5, 9, 13, and 17).
Integument moderately or sparsely setose . .
............................. 8
7. Male cerci highly bilobed, and lower apical angle
protruding more than upper (Fig. 22). Apical
lobes of male subgenital plate only slightly
ßared outward (Fig. 21). Upper carina and
upper carinula of hind femora without a row of
small dots. Two distinct dorsal bands on hind
femora with upper half of medial area black
(Fig. 23). Hind tibiae mostly red, sometime
black or blue ......S. albolineata (Thomas)
Male cerci not bilobed (Fig. 26). Apical lobes of
male subgenital plate highly ßared outward
(Fig. 25). Upper carina and upper carinula of
hind femora with a row of small dots (Fig. 27).
Two dorsal bands on hind femur present or
obliterated to no band. Hind tibiae purple,
brown or black ......S. obscura (Fabricius)
8. Fore and middle femora of males distinctively
inßated. Basal eminenceof cingulum, hour-
glass shaped, and very broad in the middle
(Figs. 41 and 45) .................. 9
Fore and middle femora of males not inßated.
Basal eminenceof cingulum, slightly or
highly constricted in the middle (Fig. 29 and
33) ..........................10
9. Overall coloration olive green. Upper carina and
upper carinula without a row of small dots
(Fig. 19). Hind tibiae mostly red, pink, or or-
ange. Posterior margin of abdominal tergites
always without a row of black dots. Found in
the southwestern United States (Arizona, Cal-
ifornia, Utah, Nevada, and Colorado) .....
................ S. shoshone (Thomas)
Coloration extremely variable (brown, olive
green, or black and yellow). Upper carina and
upper carinula with a row of small dots. Hind
tibiae brown, red, or black. Posterior margin of
abdominal tergites always with a row of black
dots. Widespread in the United States .....
...................S. lineata Scudder
10. Overall coloration rusty brown. Pronotum cy-
lindrical, not narrowing toward head. Head
slightly inßated. Dorsal longitudinal stripe
usually absent, but can be present. Male cerci
small, quadrate, length about the same as
width, distal tip slightly bilobed (Fig. 10) . .
............... S. rubiginosa (Harris)
Overall coloration brownish to slightly olive
green. Pronotum narrowing toward head.
Head small and not inßated. Dorsal longitu-
dinal stripe always present. Male cerci quad-
rate, length about the same as width, slightly
inßated in the middle, distal tip bilobed (Fig.
6)...............S. alutacea (Harris)
Schistocerca alutacea Harris, 1841 stat. rev.
(Figs. 4Ð7 and 28Ð31)
Acrydium alutaceum Harris, 1841: 139
Schistocerca alutacea (Harris) (Bruner, 1893: 26)
Schistocerca alutacea (Harris) (Henderson, 1942: 101)
Schistocerca alutacea (Harris) (Hubbell, 1960; 62)
Schistocerca alutacea alutacea (Harris) (Dirsh, 1974:
194)
Male. Medium size (total length 33Ð40 mm).
Integument moderately setose. Median carina of
pronotum distinct but not raised, with shallow sulci.
Figs. 2851. Male genitalic structures of the species in
the Alutacea Group. (28Ð31) S. alutacea; (32Ð35) S. rubigi-
nosa; (36Ð39) S. lineata; (40Ð43) S. shoshone; (44Ð47) S. al-
bolineata; (48Ð51) S. obscura; (28, 32, 36, 40, 44, and 48)
cingulum; (29, 33, 37, 41, 45, and 49) basal eminenceof
zygoma; (30, 34, 38, 42, 46, and 50) endophallus; (31, 35, 39,
43, 47, and 51) epiphallus. Ac, arch of cingulum; Anc, ancora;
Ap, apical valve of aedeagus; Apd, apodemes of cingulum; Bp,
basal valve of aedeagus; Br, bridge of epiphallus; Cv, valve of
cingulum; Gpr, gonopore process; Lp, lophus; Rm, rami of
cingulum; Zyg, zygoma. Bar a 1 mm for 28, 32, 36, 40, 44,
and 48. Bar b 1 mm for 29, 33, 37, 41, 45, and 49. Bar c
1 mm for 30, 34, 38, 42, 46, and 50. Bar d 1 mm for 31, 35,
39, 43, 47, and 51.
426 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
Hind angle of pronotum slightly angular. Lateral lobe
of metazona with granules. Pronotum slightly narrow-
ing toward head. Head small and not inßated. Fore and
middle femur not inßated. Epiproct with a pair of
tubercles (Fig. 4). Cerci quadrate, length about the
same as width, slightly inßated in the middle, distal tip
bilobed (Fig. 6). Apical lobes of subgenital plate not
outwardly ßared, notch U-shaped (Fig. 5). Phallus:
Cingulum, surfaces of rami deeply infolded in the
middle and highly convex, thus making basal emi-
nenceappear bilobate and highly constricted in the
middle (Figs. 28 and 29). Endophallus, basal valves
laterally semicircular, valves of cingulum straight,
slightly protruding more than apical valves of aedea-
gus (Fig. 30). Epiphallus, distance between lophi as
long as the length of base of a lophus (Fig. 31). Lophi
inßated triangular. Color: Rusty brown to slightly olive
green. Head yellow, rusty brown, olive green with
faint to strongly dark subocular stripes. A pair of dark
brown stripes between eyes from upper half of frontal
ridge to occiput, often extending to pronotum. Head,
pronotum, and tegmina with a yellow dorsal longitu-
dinal stripe. Lateral lobe of pronoza without marking.
Metazona sometimes with small granules. Epimeron
without marking. Tegmina brown, sometimes with
slightly mottling. Tegminal veins brown. Hind wing
with slightly yellow tinge. Posterior margin of abdom-
inal tergites with a row of black dots. Dorsal and
ventral surface of hind femur light brown. Hind femur
mostly without dorsal band, but sometimes with a
trace. Medial area white, sometimes with irregular
pattern of black dots. Carinula with a row of very small
black dots (Fig. 7). Upper carina without dots. Hind
tibia light brown, with tibial spines yellow with black
tip.
Female. Much larger than (TL 49Ð52 mm).
Median carina of pronotum slightly more raised. Oth-
erwise same as male. Ovipositor: Ventral valves long in
proÞle. Base of dorsal valves distinctively angular. Egg
guide slightly narrowing toward apex, slightly curved
upward. Pigmentation covering JannoneÕs organ dis-
tinct, sclerotized linearly in the middle.
Diagnostic Characters. Schistocerca alutacea can be
uniquely distinguished based on the highly convex
rami of cingulum (Figs. 28 and 29). It always has a
yellow dorsal longitudinal stripe. Front and middle
femora of the males are never inßated.
Material Examined. 1,223 specimens (: 726, :
497) from ANSP, CHAP, CSUC, INHS, IRCW, LEMQ,
OSEC, OSUC, SEMC, UMMZ, and UMSP.
Type Material. NEOTYPE (MCZ, here desig-
nated) S. alutacea (Harris, 1841), with labels. Mass.
MarthaÕs/Vineyard. West/Chop. Aug 1893,”“West
Chop, Mas./19 Aug 1893,”“PROPERTY/M. C. Z./
Harvard,”“TOPOTYPE/Schistocerca/alutacea (Har-
ris),”“PLESIALLOTYPE/Schistocerca/alutacea (Har-
ris),”“PLESIALLOTYPE/See Hubbell/1960: 62,
NEOTYPE/Schistocerca/alutacea (Harris)/H. Song,
2003.
Distribution. Examined specimens were collected
from Alabama, Arkansas, Connecticut, Florida, Geor-
gia, Maryland, Michigan, Mississippi, New Jersey, New
York, North Carolina, Oklahoma, Pennsylvania, South
Carolina, Tennessee, Virginia, and Wisconsin. This
species is distributed mainly in the eastern United States.
Biology. Schistocerca alutacea prefers shrubby,
moist to wet situations, including bogs, swamps,
marshes, and thickets bordering mesic forests (Hub-
bell 1960). Besides wet habitats, Squitier and Capinera
(2002a) found that the species also prefers dry sand
hill. In Florida, nymphs occur starting in May, and
adults can be frequently found in July and August
(Squitier and Capinera 2002b).
Taxonomic Discussion. Although Hubbell (1960)
considered the genitalic characters to be important for
the Alutacea Group, Dirsh (1974) apparently over-
looked these characters. His revision contained only
two drawings of the cingulum of S. alutacea alutacea,
and none for the other subspecies (Dirsh 1974). These
drawings, however, failed to show the distinct basal
eminencethat Hubbell (1960) emphasized, making it
difÞcult to validate DirshÕs taxonomic concept. I ex-
amined many phallic complexes of S. alutacea from
various collecting localities, as well as the entire gen-
italia collection used by Hubbell, and I conclude that
HubbellÕs (1960) characterization was correct, thus
reviving the taxonomic status to a valid species.
The type specimen in the collection of the Boston
Society of Natural History was destroyed (Hubbell
1960). In the original description, Harris (1841) did
not specify the sex of the type specimen, but the
original measurement (length: 1.75 in; wingspan: 3 in)
suggests that it was a female. Hubbell (1960) desig-
nated a plesiallotype, a male specimen topotypic of the
type specimen, which, however, is not recognized by
the International Code of Zoological Nomenclature
(ICZN) (International Commission of Zoological No-
menclature 2000). Vickery and Kevan (1983) consid-
ered this plesiallotype as a neotype, but no formal
action was taken. There is still some taxonomic con-
fusion about the status of S. alutacea (Capinera et al.
2001), and I think it is necessary to designate a neo-
type. Therefore, I hereby designate HubbellÕs (1960)
plesiallotype as a neotype.
Schistocerca rubiginosa Harris, 1862 stat. rev.
(Figs. 8Ð11 and 32Ð35)
Acridium rubiginosum Harris, in Scudder, 1862: 467
Schistocerca rubiginosa (Harris) (Hubbell, 1960: 66)
Schistocerca alutacea rubiginosa (Scudder) (Dirsh,
1974: 198)
Male. Medium size (TL 34Ð39 mm). Integument
without long setae. Median carina of pronotum dis-
tinct but not raised, with shallow sulci. Hind angle of
pronotum slightly angular. Pronotum without pattern
or granule. Anterior end of pronotum broad, and head
slightly inßated. Overall pronotum cylindrical. Fore
and middle femur not inßated. Epiproct with a pair of
tubercles (Fig. 8). Cerci small, quadrate, length about
the same as width, distal tip slightly bilobed (Fig. 10).
Apical lobes of subgenital plate not outwardly ßared,
notch U-shaped (Fig. 9). Phallus: Cingulum, surfaces
of rami infolded in the middle and moderately convex,
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 427
thus making basal eminenceappear slightly bilobate
and somewhat constricted in the middle (Figs. 32 and
33). Endophallus, basal valves laterally semicircular,
valves of cingulum straight, slightly protruding more
than apical valves of aedeagus (Fig. 34). Epiphallus,
distance between lophi as long as the length of base of
a lophus (Fig. 35). Lophi inßated triangular. Color:
Rusty brown. Head rusty brown with numerous small
black dots, with faintly brown subocular stripes. A pair
of rows of black dots between eyes from fastigium to
occiput. Dorsal longitudinal stripe rarely present.
Epimeron without pattern. Antennae rusty brown.
Tegmina rusty brown with slight mottling. Tegminal
veins rusty brown. Hind wing with yellow tinge. Pos-
terior margin of abdominal tergites with a row of black
dots. Dorsal and ventral surface of hind femur rusty
brown. No dorsal band on hind femur. Medial area
white to light brown with irregular pattern of black
dots (Fig. 11). Upper and lower carinula with a row of
black dots. Upper carina with a row of dots. Hind tibia
light to rusty brown, with tibial spines yellow with
black tip.
Female. Much larger than (TL 49Ð54 mm).
Median carina of pronotum slightly more raised. Oth-
erwise same as male. Ovipositor: Ventral valves long in
proÞle. Base of dorsal valves distinctively angular. Egg
guide slightly narrowing toward apex, slightly curved
upward. Pigmentation covering JannoneÕs organ dis-
tinct, faintly sclerotized linearly in the middle.
Diagnostic Characters. Schistocerca rubiginosa can
be distinguished by the slightly inßated head and the
cylindrical pronotum. It also has the smallest male
cerci (Fig. 10) compared with other species in the
group. Overall coloration is almost always rusty
brown, and most southern populations do not have a
dorsal longitudinal stripe. This species is different from S.
alutacea in the morphology of the rami of cingulum,
which is only slightly convex (Figs. 32 and 33).
Material Examined. 1,292 specimens (: 795, :
497) from ANSP, CSUC, IRCW, LEMQ, OSEC,
SEMC, UMMZ, UMSP, UNSM, and WFRM.
Type Material. NEOTYPE (UMMZ) S. rubiginosa
(Harris, 1862), with labels. BEAUFORT CO. S.C./1.1
mile N. Limehouse/(US17) 20 Aug, Õ47/2 T.H.Hub-
bell,”“PLESIALLOTYPE/Schistocerca/rubiginosa
(Harris),”“PLESIALLOTYPE/See Hubbell/1960:66,
Head and sub-/genital plate Þg./Hubbell 1960: 14,
NEOTYPE/Schistocerca/rubiginosa (Harris)/H. Song,
2003.
Distribution. Examined specimens were collected
mainly from the eastern United States including Con-
necticut, Florida, Georgia, Massachusetts, New Jersey,
North Carolina, Tennessee, and South Carolina.
Biology. Schistocerca rubiginosa prefers xeric to xe-
romesic habitats, especially sandy soil (Hubbell 1960).
In Florida, it is sympatric with S. alutacea and has a
similar seasonal phenology (unpublished data).
Taxonomic Discussion. Schistocerca rubiginosa was
considered a synonym of S. alutacea for almost 60 yr
until Hubbell (1960) revised the group because of
RehnÕs (1901, 1902) erroneous observation. On the
basis of morphological characters, I conclude that
HubbellÕs (1960) characterization was correct, thus
reviving the taxonomic status to a valid species.
As in S. alutacea, Hubbell (1960) designated a ple-
siallotype, which is not recognized by the Code (In-
ternational Commission of Zoological Nomenclature
2000). Vickery and Kevan (1983) considered this as a
neotype, but no formal action was taken. The rusty
brown color of the species has sometimes caused mis-
identiÞcation, especially with female specimens, and
even Dirsh erroneously identiÞed it as either S. dam-
nifica or S. lineata (unpublished data). Capinera et al.
(2001) and Squitier and Capinera (2002a, b) did not
recognize the species and considered it as a synonym
of S. alutacea. To prevent further taxonomic confusion,
I think it is necessary to designate a neotype. Thus, I
hereby designate HubbellÕs (1960) plesiallotype as a
neotype.
Schistocerca lineata Scudder, 1899 stat. rev.
(Figs. 12Ð15, 36Ð39, and 28Ð31)
Acridium emarginatum Uhler, manuscript
Acridium emarginatum,nomen nudum Dodge, 1872: 15
Acridium emarginatum Scudder, 1872: 250
Schistocerca lineata Scudder, 1899: 465
Schistocerca scudderi Bruner, 1906, unnecessary re-
placement name
Schistocerca lineata Scudder (Hubbell, 1960: 71)
Schistocerca emarginata (Scudder) (Vickery and
Kevan, 1964: 1555; 1983: 725)
Schistocerca alutacea lineata (Scudder) (Dirsh, 1974:
204)
Male. Medium size (TL 32Ð50 mm). Integument
sparsely setose. Median carina of pronotum distinct
and slightly raised, with distinct sulci. Hind angle of
pronotum broadly angular. Lateral lobe of metazona
mostly with granules. Pronotum slightly narrowing to
head, and head medium size. Fore and middle femur
highly inßated. Epiproct with a pair of tubercles (Fig.
12). Cerci quadrate, length about the same as width,
slightly inßated in the middle, distal tip bilobed (Fig.
14). Lower apical angle of cerci protruding slightly
more than upper. Apical lobes of subgenital plate not
outwardly ßared, notch U-shaped (Fig. 13). Phallus:
Cingulum, surfaces or rami not infolded and sinuate,
thus making basal eminenceappear hourglassÐ
shaped, and broad in the middle (Figs. 36 and 37).
Endophallus, basal valves laterally semicircular, valves
of cingulum straight, slightly protruding more than
apical valves of aedeagus (Fig. 38). Epiphallus, dis-
tance between lophi as long as the length of base of a
lophus (Fig. 39). Lophi inßated triangular. Color:
Hubbell (1960) analyzed the color variation of the
species, and here I categorize it into four distinguish-
able ecotypes: typical (Figs. 52 and 53), brown (Figs.
54 and 55), aposematic (Figs. 56 and 57), and olive
green (Figs. 58 and 59). Here I characterize the typical
form, and the differences among ecotypes are shown
in Table 3. Head light to greenish brown with brown
subocular stripes. A pair of light brown stripes be-
tween eyes from upper half of frontal ridge to occiput.
Head, pronotum, and tegmina usually with a light
428 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
yellow dorsal longitudinal stripe. Lateral lobe of pro-
zona without distinct marking. Metazona with small
light yellow granules. Epimeron with small yellow
granules. Tegmina light brown, with tegminal veins
light brown. Hind wing with slightly yellow tinge.
Posterior margin of abdominal tergites with a row of
black dots. Dorsal and ventral surface of hind femur
light brown. Hind femur without dorsal bands. Medial
area light brown. Upper and lower carinula with a row
of black dots (Fig. 15). Upper carina without dots.
Hind tibia black especially on ventral side, with tibial
spines yellow with black tip.
Female. Much larger than (TL 45Ð69 mm).
Median carina of pronotum slightly more raised. Fore
and middle femur not inßated. Otherwise same as
male. Ovipositor: Ventral valves long in proÞle. Base of
dorsal valves distinctively angular. Egg guide highly
narrowing toward apex, highly curved upward. Pig-
mentation covering JannoneÕs organ distinct, sclero-
tized linearly in the middle.
Diagnostic Characters. Schistocerca lineata is the
most polymorphic species in the Alutacea Group, and
each local population is highly variable in coloration
(Table 3) and perhaps host preference. Therefore, it
is difÞcult to characterize this species except by the
phallic morphology, where the basal eminencere-
sembles an hour-glass and is broad in the middle (Figs.
36 and 37). This genitalic character was Þrst intro-
duced by Hubbell (1960). The species also possesses
the inßated front and middle femora in males, which
are also found in S. shoshone. However, S. lineata can
Figs. 5259. Four ecotypes of S. lineata. (52Ð53) Typical
form; (54Ð55) brown form; (56Ð57) aposematic form; (58Ð
59) olive green form. Bar 1 cm.
Table 3. Four ecotypes of S. lineata and their color variation
Ecotypes Distribution Head Dorsal stripe Epimeron Tegmina Hind femur Hind tibia
Typical Great Plains
region
Light to greenish brown
with brown subocular
stripes
Light yellow With small yellow
granules
Light brown with light
brown veins
Light brown without dorsal bands;
carinula with a row of black
dots; upper carina without dots
Black especially on ventral side,
with tibial spines yellow with
black tip
Brown Great Lake
regions
Brown with faint
subocular stripes
Absent Without pattern Light brown with
slight mottling, with
brown veins
Brown without dorsal bands;
carinula with a row of black
dots; upper carina sometimes
with a row of dots
Light brown, with tibial spines
yellow with black tip
Aposematic Southwestern U.S. Yellow with black
subocular stripes
Yellow Entirely yellow
without distinct
marking, but
sulci between
episternum and
epimeron black.
Yellowish brown,
sometimes with
slight mottling, with
brown veins
Light brown to yellow with
distinct black dorsal bands;
carinula with/without a row of
very small black dots; upper
carina mostly without dots
Black especially on ventral side,
with tibial spines yellow with
black tip
Olive green Western United
States
Olive green with dark
green subocular
stripes
Bright yellow With small yellow
granules
Light brown to olive
green, with lemon
yellow veins
Olive green sometimes with two
dorsal bands; carinula
sometimes with a row of black
dots, but usually absent; upper
carina sometimes with dots, but
usually absent.
Pinkish red, with tibial spines
yellow with black tip
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 429
always be distinguished from S. shoshone by the pres-
ence of the black dots on the abdominal tergites.
Material Examined. 3,151 specimens (: 1656, :
1495) from ANSP, BYU, CSUC, INHS, IRCW, LEMQ,
NMSU, OSEC, OSUC, SEMC, UARM, UMMZ, UMSP,
UNSM, and WFRM.
Type Material. LECTOTYPE (ANSP) S. lineata
Scudder, 1899, with labels. Barbour Co./Kas. Cragin,
Schist./lineata/ScudderÕs/Type, 1899,”“Ex Coll.L.
Bruner/S. lineata Sc./Proc. Am. Ac. A.&S./Vol.
XXXIV p. 466./Hebard Collection,”“Schistocerca/lin-
eata Sc./TYPE H86,”“Sch. alutacea/lineata Sc./Lect./
V.M.Dirsh det., 1972.
Distribution. This is the most widely distributed
species of Schistocerca in North America. Examined
specimens were collected from Arizona, Arkansas,
Connecticut, Colorado, Delaware, Georgia, Indiana,
Illinois, Idaho, Iowa, Kansas, Kentucky, Massachu-
setts, Maryland, Michigan, Minnesota, Mississippi, Ne-
braska, New Jersey, New Hampshire, New Mexico,
New York, North Dakota, Ohio, Oklahoma, Rhode
Island, Sout Dakota, Texas, Utah, Virginia, and Wis-
consin. Its distribution extends north to Manitoba and
Alberta, Canada.
Biology. Schistocerca lineata is abundant in sandy
areas but is also frequently found in other habitats.
Detailed account of its habitat association is found in
Hubbell (1960). Criddle (1932) reported that the de-
velopment takes 39 d from the time of hatching. He
also noted that this species has Þve nymphal instars.
Sword et al. have shown that the Texas population has
a density-dependent polyphenism in nymphal instar,
mediated by the host preference (Sword 1999, Sword
and Dopman 1999, Dopman et al. 2002, Sword 2002).
Taxonomic Discussion. Here, I designate four
ecotypes of S. lineata. Dopman et al. (2002) found that
the populations associated with either Ptelea or Rubus
form a monophyletic clade using 16S rRNA and 12S
rRNA regions of the mitochondrial DNA. Although
their study demonstrated the lack of gene ßow be-
tween two host-associated populations, I am hesitant
to create a new taxonomic concept at this moment,
because not much is known about the populations in
the midwestern and eastern United States. Highly con-
spicuous individuals from the southwestern states are
more robust in form and slightly larger than noncon-
spicuous ones. In terms of coloration, specimens from
Great Lake regions are brown and resemble S. rubigi-
nosa. Aposematic specimens are never found in north-
ern states, although the polymorphism of dorsal lon-
gitudinal stripe exists. In western states, specimens
become almost indistinguishable from S. shoshone ex-
cept on the basis of markings on abdominal tergites.
These specimens have olive green hue, which is rarely
observed in either southwestern or eastern popula-
tions. In all cases, however, crucial morphological
characters, such as phallic complex and inßated male
femur, are identical.
The main reason Dirsh (1974) changed the ranks of
previously recognized species in the Alutacea Group
was his claim to have found the hybrid between S.
lineata and S. albolineata. I have examined several
specimens bearing labels by Dirsh (1974) stating,
Schistocerca alutacea lineata transient to albolin-
eata.These specimens are, however, not hybrids, but
aposematic individuals of S. lineata found in Texas. S.
lineata is different from S. albolineata in terms of the
phallic complex (Figs. 36Ð39) and the front and mid-
dle femora in males. Therefore, I argue that DirshÕs
(1974) synonymy was based on the incorrect assump-
tions about the biology of the species, and here I revive
the taxonomic status to a valid species.
There is also a long-standing confusion on the va-
lidity of the name S. lineata. Detailed account can be
found in Appendix C, and here I argue that S. lineata
is a valid name as nomen protectum.
Schistocerca shoshone Thomas, 1873 stat. rev.
(Figs. 16Ð19 and 40Ð43)
Acridium shoshone Thomas, 1873: 295
Schistocerca venusta Scudder, 1899: 467 (Hebard, 1935:
299)
Schistocerca obliquata Scudder, 1899: 470 (Hebard,
1932: 279)
Schistocerca shoshone (Thomas) (Henderson, 1942:
99)
Schistocerca alutacea shoshone (Thomas) (Dirsh, 1974:
200)
Male. Medium size (TL 39Ð44 mm). Integument
without long setae. Median carina of pronotum dis-
tinct and slightly raised, with distinct sulci. Metazona
with numerous small granules. Hind angle of prono-
tum slightly angular. Fore and middle femur highly
inßated. Epiproct with a pair of tubercles (Fig. 16).
Cerci quadrate, length about the same as width,
slightly inßated in the middle, distal tip bilobed (Fig.
18). Apical lobes of subgenital plate not outwardly
ßared, notch U-shaped (Fig. 17). Phallus: Cingulum,
surfaces or rami not infolded and sinuate, thus making
basal eminenceappear hourglass-shaped and very
broad in the middle (Figs. 40 and 41). Endophallus,
basal valves laterally semicircular, valves of cingulum
straight, slightly protruding more than apical valves of
aedeagus (Fig. 42). Epiphallus, distance between lo-
phi as long as the length of base of a lophus (Fig. 43).
Lophi inßated triangular. Color: Mostly light olive
green, occasionally lemon yellow or slight brown. Two
forms can be distinguished based on the presence of
dorsal stripe. Unstriped form: Head olive green to yel-
low with dark olive green to brown subocular stripe.
No pair of stripes between eyes, and no dorsal longi-
tudinal stripe on head and pronotum. Lateral lobe of
pronoza without marking. Metazona sometimes with
small granules. Antennae yellow. No pattern on
epimeron. Tegmina light olive green to yellow without
any pattern. Tegminal veins light lemon-yellow to
brown. Hind wing with slightly yellow tinge. Posterior
margin of abdominal tergites without a row of black
dots. Dorsal and ventral hind femur yellow to olive
green. Medial area white without any pattern. Hind
femur without dorsal bands. Carinula without pattern
(Fig. 54). Hind tibia pink, orange to red, with tibial
spines yellow with black tip. Striped form: Head light
430 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
olive green to yellow with faint dark olive green to
brown subocular stripes. A pair of olive green stripes
between eyes from upper half of frontal ridge to oc-
ciput. Head, pronotum, and tegmina with a yellow
dorsal longitudinal stripe. Lateral lobe of prozona with
no apparent marking, and lower half of prozona and
entire metazona with yellow granules. Second
epimeron sometimes with yellow marking. Antennae,
tegmina, hind wing, abdomen, and hind femur same as
unstriped form.
Female. Much larger than (TL 53Ð63 mm).
Median carina of pronotum distinctly more raised.
Fore and middle femur not inßated. Otherwise same
as male. Ovipositor: Ventral valves short in proÞle.
Base of dorsal valves distinctively angular. Egg guide
highly narrowing toward apex, highly curved upward.
Pigmentation covering JannoneÕs organ distinct, scle-
rotized linearly at the base.
Diagnostic Characters. Schistocerca shoshone can be
easily distinguished by the absence of black markings
on the abdominal tergites. These black markings ap-
pear to be the universal color pattern in Schistocerca
species, but S. shoshone always lacks this pattern. This
species is closely related to S. lineata, because both
share the similar phallic morphology and inßated fore
and middle femora in males.
Material Examined. 991 specimens (: 498, : 493)
from ANSP, BYU, CSUC, INHS, IRCW, LEMQ, OSEC,
OSUC, NMSU, SEMC, UARM, UMMZ, UMSP, UNSM,
and WFRM.
Type Material. 1. NEOTYPE lost. Utah, Cache
County, Logan(Dirsh 1965) I was unable to Þnd the
neotype at ANSP. 2. LECTOTYPE (ANSP, dis-
sected phallus stored in a vial) S. venusta Scudder,
1899, with labels. Indio, Calif./9 July, 1897,”“Schist./
venusta/ScudderÕs/Type, 1899,”“S.H.Scudder/Coll.,
Type/1829,”“Lecto/type/R.&H. 1912,”“venusta/
Scudder.3. LECTOTYPE (ANSP, dissected phallus
stored in a vial) S. obliquata Scudder, 1899, with labels.
San Jose/del cabo/Mexico,”“Schist./obliquata/Scud-
derÕs/Type, 1899,”“Ex Coll.L.Bruner/S. obliquata Sc./
Proc. Am. Ac. A.&S./Vol. XXXIV p. 471/Hebard Col-
lection,”“Schistocerca/obliquata Sc./TYPE H393,
Lectotype.
Distribution. Schistocerca shoshone occurs in the
western United States including Arizona, California,
Colorado, Idaho, Nevada, New Mexico, Oregon,
Texas, and Utah.
Biology. It is present in both riparian and desert
habitats and prefers various woody plants (Sword and
Chapman 1994). In Utah, it is frequently found in
cornÞelds or other tall growing vegetation (Hender-
son 1942). This species is univoltine, and adults occur
in June and July (Chapman et al. 1995).
Taxonomic Discussion. Schistocerca shoshone has
both striped and unstriped forms that seem to be
associated with certain host plants (Chapman et al.
1995). Because these two forms can apparently inter-
breed (Chapman and Sword, unpublished data), I
hereby call these ecotypes of a single species.
DirshÕs (1974) redescription of S. alutacea shoshone
is suspicious, because it states that the anterior femora
of males are slightly or not at all inßated. All type
specimens as well as all the examined specimens pos-
sess this character, and usually the inßation is even
more obvious than in S. lineata. Therefore, I argue that
DirshÕs synonymy is erroneous, and I hereby revive
the taxonomic status to a valid species.
Schistocerca albolineata Thomas, 1875 stat. rev., syn.
nov.
(Figs. 20Ð23 and 44Ð47)
Acridium albolineata Thomas, 1875: 897
Schistocerca mexicana Scudder, 1899: 468 (Hebard,
1932: 281)
Schistocerca insignis Hebard 1932: 279 (syn. nov.)
Schistocerca chinatiensis Tinkham 1948: 607 (Dirsh,
1974: 209)
Schistocerca alutacea insignis (Hebard) (Dirsh, 1974:
212)
Schistocerca alutacea albolineata (Thomas) (Dirsh,
1974: 209)
Male. Medium size (TL 39Ð43 mm). Integument
setose, with long setae on pronotum, thorax, cerci, and
subgenital plate. Median carina slightly raised, with
distinct sulci. Metazona with numerous small granules.
Hind angle of pronotum slightly angular. Width of
metazona slightly wider than width of prozona. Fore
and middle femur not inßated. A pair of tubercles on
epiproct absent (Fig. 20). Cerci quadrate, length 1.5
times longer than width, and bilobed (Fig. 22). Lower
apical angle of cerci protruding more than upper.
Apical lobes of subgenital plate slightly ßared out-
ward, notch round V-shaped (Fig. 21). Phallus: Cin-
gulum, surfaces of rami deeply infolded in the middle
and highly convex, thus making basal eminenceap-
pear slightly bilobate and somewhat constricted in the
middle (Figs. 44 and 45). Endophallus, basal valves
ventral angle protruding more than dorsal, valves of
cingulum club-shaped and curved downward, pro-
truding more than apical valves of aedeagus (Fig. 46).
Epiphallus, distance between lophi longer than the
length of base of a lophus (Fig. 47). Lophi lamelliform.
Color: Olive brown. Head yellowish to olive brown
with dark-brown to black subocular stripes. A pair of
olive brown stripes between eyes from upper half of
frontal ridge to occiput. Head and pronotum with a
bright yellow dorsal longitudinal stripe, sometimes
extending to tegmina. Epimeron often distinctly yel-
low. Antennae yellow. Lateral lobe of pronotum some-
times with a yellow rectangular spot. Tegmina uni-
formly olive brown, with veins light-brown to yellow.
Hind wing with yellow tinge. Posterior margin of ab-
dominal tergites with a row of black dots. Dorsal and
ventral surface of hind femur yellow, with two distinct
black dorsal bands. Posterior tip of hind femur black.
Outer face of hind femur, upper half of medial area
sometimes black and lower half white (Fig. 23). Ca-
rinula and carina without spots. Hind tibia mostly red
or orange-red, but occasionally black or blue, with
tibial spines yellow with black tip.
Female. Much larger than (TL 49Ð61 mm).
Median carina of pronotum more raised. Otherwise
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 431
same as male. Ovipositor: Ventral valves short in pro-
Þle. Base of dorsal valves not angular. Egg guide not
narrowing toward apex, slightly curved upward. Pig-
mentation covering JannoneÕs organ distinct, sclero-
tized circularly in the middle.
Diagnostic Characters. Males of S. albolineata pos-
ses elongated cerci with the lower apical angle pro-
truding more than upper (Fig. 22), which is unique
among the species in the Alutacea Group. The fol-
lowing characters can also be found in S. obscura, but
the two species are never sympatric. They also have an
elongated subgenital plate that is slightly ßared out-
ward (Fig. 21). The epiproct lacks a pair of tubercles
(Fig. 20). The epiphallus is wide in the bridge between
lophi (Fig. 47), and the apical valve of cingulum is
clubbed (Fig. 46). Most known specimens have a
bright yellow dorsal longitudinal stripe, two black dor-
sal bands on hind femur, and red hind tibia.
Material Examined. 240 specimens (: 140, : 100)
from ANSP, BYU, NMSU, and UMMZ.
Type Material. 1. NEOTYPE (ANSP) S. albolin-
eata Thomas, 1875, with labels. Ajo, Pima Co. Ariz./
ab.1800 feet (R&H)/IX, 18, 1922,”“Schistocerca/
alutacea albolineata Thom./V.M. Dirsh det. 1971,
NEO-TYPE.2. HOLOTYPE (ANSP, dissected
phallus stored in a vial) S. mexicana Scudder, 1899,
with labels. Sinaola, Mex./KOELS. J. BEHRENS,
Schist,/mexicana/ScudderÕs/Type, 1899,”“S.H.Scud-
der/Coll.,”“Type 1830,”“Holotype,”“HOLOTYPE/
Sch. mexicana/Scudder, 1899./V.M. Dirsh 1964.3.
HOLOTYPE (ANSP, dissected phallus stored in a
vial) S. insignis Hebard, 1932, with labels. Guadalajara
Jalisco/9.19 1903. Mex./J.F. McClendon,”“Schisto-
cerca/insignis/Hebard,/TYPE 5509.4. ALLOTYPE
(ANSP) S. insignis Hebard, 1932, with labels.
Guadalajara Jalisco/9.18 1903. Mex./J.F. McClen-
don,”“Schistocerca/insignis/Hebard,/Allotype .5.
HOLOTYPE (CAS) S. chinatiensis Tinkham 1948,
with labels. Chinatis/3 miles W of/Schafter/19 Oct
46,”“Type/Schistocerca/chinatiensis/msp. 1947
Tink.,”“ERNEST ROBERT TINKHAM/COLLEC-
TIONÑ1988/DONATED TO THE CALIFORNIA/
ACADEMY OF SCIENCES/by Marion Blair
Tinkham,”“California Academy/of Sciences/Type no.
16968.
Distribution. Schistocerca albolineata occurs in the
southwestern United States including Arizona, New
Mexico, and Texas. Its distribution extends south to
Jalisco, Mexico.
Biology. Schistocerca albolineata prefers dry moun-
tainous habitats. It usually feeds on various woody
plants, and adults can be found in early September
(unpublished data). On Tucson Mountain, AZ, I found
a population of emerging adults feeding on foothill
Palo Verde Cercidium microphyllum (Torrey) Rose
and Johnston (Fabaceae). Howard (1995) found that
the Arizona populations preferred cotton Gossypium
hirsutum L. (Malvaceae).
Taxonomic Discussion. Although S. albolineata is
the most homogeneous species in the Alutacea Group
in terms of coloration, examination of a large number
of specimens revealed that the color variation indeed
exists especially in hind tibia. Color variation in Schis-
tocerca is well known (Hubbell 1960), and there seems
to be the southward darkening progression in the
species. S. chinatiensis Tinkham, described based on a
local population from the Chinati Mountains, TX, has
slightly darker coloration and black tibia. S. insignis
Hebard, only known from eight specimens from
Guadalajara, Mexico, is even darker in general colora-
tion and has black tibia. Hubbell (1960) suggested that
it is possible for S. albolineata, S. chinatiensis, and S.
insignis to be of the same polytypic species, and I agree
with his idea. All known specimens of the populations
from Arizona, New Mexico, Texas, and Mexico possess
the invariable diagnostic characters. Therefore, I ac-
cept DirshÕs (1974) synonymy of S. chinatiensis, and I
hereby synonymize S. insignis under S. albolineata on
the basis of morphological similarity. I also argue that
DirshÕs synonymy overlooked the distinct morpholog-
ical characters in S. albolineata, and I hereby revive the
taxonomic status to a valid species.
Schistocerca obscura Fabricius, 1798
(Figs. 24Ð27 and 48Ð51)
Gryllus obscurus Fabricius, 1798: 194
Schistocerca obscura (Fabricius) (Scudder, 1899: 465)
Schistocerca obscura (Fabricius) (Dirsh, 1974: 181)
Male. Medium size (TL 38Ð46 mm). Integument
highly setose. Median carina of pronotum distinct and
slightly raised, with distinct sulci. Hind angle of prono-
tum slightly angular. Width of metazona slightly wider
than width of prozona. Fore and middle femur not
inßated. A pair of tubercles on epiproct absent (Fig.
24). Cerci quadrate, length twice longer than width,
not bilobed (Fig. 26). Apical lobes of subgenital plate
strongly ßared outward, notch deep (Fig. 25). Phallus:
Cingulum, surfaces of rami deeply infolded in the
middle and highly convex, thus making basal emi-
nenceappear bilobate and highly constricted in the
middle (Figs. 48 and 49). Endophallus, basal valves
ventral angle protruding more than dorsal, valves of
cingulum club-shaped and curved downward, pro-
truding more than apical valves of aedeagus (Fig. 50).
Epiphallus, distance between lophi much longer than
the length of base of a lophus (Fig. 51). Lophi lamel-
liform. Color: Olive green to liver color. Head yellow-
ish to liver color with dark-brown to black subocular
stripes. A pair of dark brown stripes between eyes
from upper half of frontal ridge to occiput. Head and
pronotum with a bright yellow dorsal longitudinal
stripe, often extending to tegmina. Upper half of
epimeron often distinctly yellow. Antennae yellow to
orange. Lateral lobe of pronotum without pattern.
Lower half of lateral prozona and entire metazona
with yellow granules. Tegmina uniformly purplish
liver color except for the extending dorsal longitudinal
stripe. Tegminal veins same color as tegmina. Hind
wing transparent with yellow tinge. Posterior margin
of abdominal tergites with a row of black dots. Dorsal
and ventral surface of hind femur yellowish olive to
liver color, with two black dorsal bands which some-
times reduced to a trace. Medial area mostly white.
432 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
Upper and lower carinula on outer face with a row of
small black dots (Fig. 27). Hind tibia brown to black
with tibial spines yellow with black tip.
Female. Much larger than (TL 52Ð70 mm).
Median carina of pronotum more raised. Otherwise
same as male. Ovipositor: Ventral valves long in pro-
Þle. Base of dorsal valves distinctively angular. Egg
guide slightly narrowing toward apex, ventral portion
highly sinuate. Pigmentation covering JannoneÕs or-
gan distinct, sclerotized circularly in the middle and
patterns present on top.
Diagnostic Characters. Schistocerca obscura can be
easily distinguished from other species in the Alutacea
Group on the basis of the highly ßared male subgenital
plate (Fig. 25). Male cerci are elongated with length
twice longer than width, and not bilobed (Fig. 26). It
also has the epiproct lacking a pair of tubercles (Fig.
24), the wide epiphallus (Fig. 51), and the clubbed
apical valve of cingulum (Fig. 50), which it shares with
S. albolineata.
Material Examined. 985 specimens (: 495, : 490)
from ANSP, BYU, CHAP, CSUC, INHS, IRCW,
LEMQ, NMSU, OSEC, OSUC, SEMC, UARM, UMMZ,
UMSP, UNSM, and WFRM.
Type Material. 1. NEOTYPE (BMNH) S. obscura
(Fabricius, 1798), with labels, Allen Co Ks/Ele.
062,15/R.H. Beamer,”“NEO-/TYPE,”“Brit.Mus./
1925Ð208,”“Schistocerca/obscura/(Fabr),”“Schisto-
cerca/obscura (Fabr, 1798)/V.M. Dirsh det., 1974.
Distribution. Examined specimens were collected
from Alabama, Arkansas, Florida, Georgia, Kansas,
Louisiana, Maryland, Mississippi, Missouri, Nebraska,
New Mexico, North Carolina, Oklahoma, South Caro-
lina, Tennessee, Texas, and Virginia. Its distribution
extends south to Tamaulipas, Mexico.
Biology. Duck (1944) documented the bionomics of
S. obscura after the grasshopper outbreak in Okla-
homa. It prefers to feed on woody plants such as
American elm and cotton. The species has Þve
nymphal instars and emerges as an adult 50 d after
hatching. Populations in central and south Texas are
commonly associated with hackberry, Celtis laevigata
Willd. (Ulmaceae) (Chapman and Sword, unpub-
lished data). Its preferred habitat is Þelds and open
woodlands (Capinera et al. 2001). Emerging adults
were collected in August in Florida (unpublished
data).
Taxonomic Discussion. Dirsh (1974) treated S. ob-
scura as a species distinct from S. alutacea based on the
shape of subgenital plate (Fig. 25). However, the
present phylogeny suggests that S. obscura is sister to
S. albolineata indicating the paraphyly of S. alutacea
sensu Dirsh (1974). No taxonomic action is taken for
S. obscura in this revision, but this species is now
considered to be included in the Alutacea Group.
Acknowledgments
I thank J. Wenzel for continuous guidance and N. Johnson
for offering expertise on taxonomy. I am grateful to the
curators of the museums from where I borrowed specimens,
especially to D. Otte and J. Weintraub (ANSP) and L.
Knowles and M. OÕBrien (UMMZ) for allowing me to work
at their collections. P. Perkins (MCZ) searched for the miss-
ing type. I thank G. Sword (USDA) for guidance and for
sharing unpublished data and T. Cohn (UMMZ) for stimu-
lating discussion on the nomenclature. J. Capinera (Univer-
sity of Florida) and the late R. Chapman (University of
Arizona) provided invaluable information on the ecology of
the grasshoppers during the Þeldwork. I thank the members
of Wenzel Lab, K. Pickett, S. Bucheli, and J. Raczkowski for
reviewing the manuscript. I also thank T. Cohn and two
anonymous reviewers for valuable comments on the manu-
script. This work was funded by the National Science Foun-
dation Graduate Research Fellowship and partially by the
OrthopteristsÕ Society Research Grant.
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Appendix A: Characters and Character States
0. Antennae length: as long as head pronotum
0; longer than head pronotum 1; shorter than
head pronotum 2.
1. Antennae length (from 0:1): slightly longer than
head pronotum 0; much longer than head
pronotum 1.
2. Tegmina length: much longer than tip of abdo-
men 0; slightly longer than tip of abdomen 1;
highly reduced 2.
3. Distal end of tegmina color: partially transpar-
ent 0; nontransparent 1.
4. Tegmina pattern: no pattern or weak mottling
0; distinct mottling 1; numerous dark patches
2; distinct bands 3.
5. Outer margin of tegmina: straight 0; curved
1.
6. Male subgenital plate: subconical 0; weakly tri-
lobate 1; bilobate 2.
7. Shape of apical lobes of male subgenital plate
(from 6:2): pointed 0; round not ßared out-
ward 1; round and ßared outward 2.
8. Width of apex of male subgenital plate (from 6:2):
apex as long as base 0; apex slightly narrower
than base 1; apex much narrower than base
2.
9. Male cerci: conical 0; quadrate 1.
10. Quadrate male cerci (from 9:1): tapering toward
apex 0; not tapering toward apex 1.
11. Male supra anal plate: a pair of tubercles absent
0; a pair of tubercles present 1.
12. Inßated male fore- and middle femur: absent
0; present 1.
13. Posterior margin of metazona: round 0; semi-
pointed 1; highly pointed 2.
14. Pronotum setae: sparse 0; numerous 1.
15. Pronotum with small granules: absent 0;
present 1. .
16. Prosternal process: curved backward and reach-
ing sternum 0; straight or only slightly
curved 1.
17. Apical valve of aedeagus: very long 0; short
1.
18. Endophallus (from 17:1): valve of cingulum
pointed 0; valve of cingulum clubbed 1.
19. Epiphallus: bridge very long 0; bridge short
1.
20. Epiphallus (from 19:1): bridge shorter than the
base of lophus 0; bridge as long as the base of
lophus 1; bridge slightly longer than the base
of lophus 2.
21. Cingulum: elongated 0; short 1.
22. Basal eminence of cingulum (from 21:1): not
constricted 0; constricted 1.
Appendix B: Character Matrix
Appendix C: Case for the Valid Taxonomic Usage of
S. lineata
Schistocerca lineata has sometimes been called S.
emarginata by a few authors (Vickery and Kevan 1964,
1967, 1983, Cantrall 1968, Sword and Dopman 1999,
Sword 2001, Dopman et al. 2002, Sword 2002,) because
of an invalid action by Vickery and Kevan (1964) to
strictly adhere to the Principle of Priority. The taxo-
nomic history of S. lineata is rather complex and is
partially summarized in Hubbell (1960) and Vickery
and Kevan (1964, 1983). Here, I explain how confusion
arose and argue why S. lineata is the valid name.
Uhler characterized Acridium emarginatum only in
a manuscript and never published it, and Dodge
(1871) was the Þrst one to use this name as nomen
nudum when he listed several grasshopper species
collected in Nebraska. Hayden (1872) characterized
A. emarginatum and attributed this name to Uhler but
noted that the name came only from the manuscript.
Through this paper, Hayden (1872) inadvertently es-
tablished himself as author of A. emarginatum. The
type specimen of A. emarginatum was lost, and a neo-
type was never designated (Hubbell 1960). In 1899,
Scudder (1899) incorrectly synonymized A. emar-
ginatum with S. alutacea and described a new species,
S. lineata. From 1899Ð1960, S. lineata was used con-
tinuously and unambiguously with two exceptions
(Kellogg 1905, Osborn 1939). Kellogg (1905) used S.
emarginata under a Þgure copied from Lugger (1898),
and the Þgure alone is difÞcult to associate with the
name because many Schistocerca species are externally
similar. Osborn (1939) merely listed several names of
injurious grasshoppers, and the name alone is difÞcult
to associate with a taxonomic concept. In 1960, Hub-
bell (1960) published a thorough taxonomic revision
on S. lineata, S. alutacea and S. rubiginosa. He acknowl-
edged that lineata is a strict synonym of emarginata but
nevertheless retained S. lineata because of the pre-
vailing use. His revision effectively reversed the pre-
cedence under Articles 23.9 of the ICZN, and S. lineata
became a valid name as nomen protectum. In 1964,
Vickery and Kevan (1964) reinstated S. emarginata
based on the Principle of Priority. Their justiÞcation
0 5 10 15 20
兩兩兩兩兩
Ornithacris 0-00300--0-0020000-0-0-
Cyrtacanthacris 0-00200--0-0020000-0-0-
Acanthacris 0-11200--0-0020000-0-0-
Halmenus 0-2--01--0-000001101110
S. americana 0-002020110000001101010
S. nitens 10001020210001001101110
S. damnifica 2-110120210002101101210
S. ceratiola 11101120110101101101110
S. alutacea 11110121111101111101111
S. rubiginosa 11110121111101111101111
S. lineata 11110121111111111101110
S. shoshone 11110121111111111101110
S. albolineata 11110122011001111111211
S. obscura 11110122011001111111211
May 2004 SONG:REVISION OF THE ALUTACEA GROUP OF Schistocerca 435
for this action came from the prevailing usage of the
name between 1872 and 1898. However, A. emar-
ginatum was the only available name during that pe-
riod, and S. lineata was described in 1899. Because the
name emarginata was already suppressed by Hubbell
(1960) in accordance with the Code (International
Commission of Zoological Nomenclature 2000), Vick-
ery and Kevan (1964) could not have simply reinstated
the name. To do so, they should have submitted a
proposal to the Commission, which they never did.
Between 1964 and 1974, S. emarginata was used once
by Vickery and Kevan (1967). In the complete revi-
sion of the genus, Dirsh (1974) agreed with HubbellÕs
(1960) use of lineata, even though his discussion of
synonymy indicated that emarginata has priority. Al-
though he cited Vickery and Kevan (1964), Dirsh
(1974) did not mention their action in the discussion
perhaps because he rejected their proposal. Vickery
and Kevan (1983), apparently troubled by DirshÕs
revision, dismissed the revision in its entirety. Once
again, they reinstated S. emarginata. Since then, S.
emarginata (lineata) was used in a series of ecological
studies by Sword and Dopman (1999), Sword (2001),
Dopman et al. (2002), and Sword (2002).
It is true that the name A. emarginatum (or S. emar-
ginata) is older, thus taking precedence to S. lineata.
HaydenÕs (1872) description is sufÞcient to make him
an author but insufÞcient to distinguish A. emar-
ginatum from other Schistocerca species. With inade-
quate characterization and no type specimen avail-
able, the name itself has a dubious status. Hubbell
(1960) discovered this older name but nevertheless
used S. lineata in his revision because of the prevailing
use of the latter name. Because the name emarginatum
had not been used in the primary zoological literature
for 50 yr, and because the later name lineata was
being used prevalently, Hubbell (1960) did not con-
sider it necessary to keep the antiquated name emar-
ginatum. To reverse the precedence based on the 50-yr
rule, one needs to apply to the Commission only if it
is proposed from 1961. Interestingly, HubbellÕs paper
was published on 29 December 1960. Therefore, one
can assume that, under the rules in existence at that
time, Hubbell (1960) did not need the action of the
Commission to declare a name as nomen oblitum. Hub-
bell (1960) appreciated the objectives of the Code,
which are to promote stability and not upset a long-
accepted name in its accustomed meaning. Vickery
and KevanÕs (1964) subsequent actions were errone-
ous and irrelevant. Indeed, resurrecting an obscure
name has a negative effect in maintaining stability of
taxonomy. Cohn (1999) criticized the use of an ob-
scure name that takes precedence to a widely used
later name, using an example of Romalea microptera
(Beauvois, 1817). Kevan (1980) discovered an older
name of this common Eastern lubber grasshopper and
argued for its name change to Romalea guttata (Stoll,
1813) on the basis of Principle of Priority. This name
had not been used for 150 yr, and it should have been
suppressed because introduction of this name would
endanger the stability of already well-established use
of R. microptera. As a result, both names were used to
refer to one taxonomic concept since 1980, which is,
of course, far from the stability that ICZN strives for.
Similar confusion has already happened in Schistocerca
where some researchers used S. emarginata (Sword
and Dopman 1999, Sword 2001, Dopman et al. 2002,
and Sword 2002) and others used S. lineata (Otte 1970,
Helfer 1972, Capinera and Sechrist 1982, Richman et
al. 1993, Hubbell 1960) to refer to the same taxonomic
concept. To maintain the stability and universality of
a taxonomic name, partially hindered by Vickery and
KevanÕs (1964) action, I argue that HubbellÕs (1960)
initial action was correct, effectively making S. lineata
a valid name as nomen protectum.
436 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 3
... Schistocerca nitens conocida comúnmente como "saltamontes gris" ("Gray bird grasshopper"), es una especie de acrídido que generalmente se le considera como solitaria y no migratoria, pero que bajo ciertas condiciones climáticas puede transformarse en gregaria y formar enjambres o brotes invasivos que causan daños a los cultivos y flora nativa; la misma posee un amplio rango de distribución geográfica nativa o autóctona, desde los EUA hasta Brasil (Song 2004, Cigliano et al. 2021 Como en muchos taxones del género Schistocerca, el estatus taxonómico de S. nitens aún no se encuentra establecido adecuadamente, por lo que se le considera como la especie que tiene mayores problemas taxonómicos dentro del género. Esto aparece debido en primera instancia, a que por su amplio rango de distribución geográfica posee poblaciones adaptadas a los hábitats locales, generando una amplia variabilidad morfológica, además de que tiene una estrecha semejanza morfológica con especies afines. ...
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... 4236/gep.2020.812009 ical African species considered as the most notorious insects in the world and has the largest geographical distribution spreading over 30 countries (Wilps, 1997;Song, 2004). The peculiarity of Schistocerca gregaria is that it exhibits phenotypic plasticity called density-dependent phase polyphenism in which local population density affects the expression of a variety of behavioral, physiological, biochemical, molecular, reproductive, and morphological traits (Cabej, 2019). ...
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