THE INTERNAI, MALE GENITALIA OF SELECTED
GENERA OF MELANOPLINAE (ORTHOPTERA: ACRIDIDAE).
by
Spyros D. Skareas
Depa rtment of Na tural Resource Sciences
McGill University, Montreal
June 1998
A thesis subrnitted to the Faculty of Graduate Studies and Research in partial
fulfillment of the requirements of the degree of Master of Science
0 Spyros D. Skartas
1998
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ABSTRACT
The internai male genitdia of selected genera of Melanoplinae (Onhoptera: Acndidae).
M. Sc. Spyros D. Skareas Natural Resource Sciences
The morphology of the intemal male genitaiia of selected genera of Melanoplinae
(Onhoptera: Acrididae) was studied. A detailed cornparison of these structures was
carried out, in an attempt to identiQ new characters that may be usehl in analyzing
relationships between the members of the subfamily. Twenty two genera were examined,
mostly of North American distribution, using one exernplar species for each of them.
Standard dissection and drawing techniques were followed. The epiphallus and the apical
parts of the aedeagus were found to be the most important taxonomie characters. The
genera Apfenopedes and Buckeiîacris were highly divergent from the rest of the
melanoplines, while the close relationship between Appalachia, Dendrotettix and Podisma
was confirmed by genital characters. Internai male genitalic characters did not provide
. significant support for many currendy recognized tribes and subtribes, suggesting that
there is a great need for reclassification in the subfarnily.
F~ÉSUME
Les structures génitales mâles des genres sélectionnés de la sous-famille
Mélanoplinae (Orthoptera: Acrididae)
M. Sc. Spyros D. Skareas Sciences des Ressources Naturelles
La morphologie des genitalia m l e des genres sélectionnés de la sous-famille
Mélanoplinae a été réalisée, afin de permettre d'identifier de nouveaux charactères qui
pourraient être utiles dans l'analyse des liens entre les membres de cette sous-famille. Une
espèce pour chacun des vingt-deux genres a été étudiée, majoritairement de provenance
nord-américaine. Des techniques de dissection et de dessin coumantes ont été utilisées. II
a été conclu que I'épiphaile et les parties apicales de l'édéage sont les charactères
taxonomiques les plus importants. Les genres Aptenopedes et Brtckellacris étaient très
divergents du reste des Mélanoplines. Le lien étroit entre Appalachia, Dendroteztix et
Pudisma a été confirmé par l'examen des charactères génitaux. Les structures génitales
mâles ne fournissent pas d'appui significatif pour plusieurs tribus et sous-tribus reconnues,
indiquant qu'il est nécessaire de reclassifier la sous-famille.
To Dr. Mazomenos & D r Hsiung for introducing me to
behavioral and systematic Entomology
AKNOWLEDGEMENTS
1 would like to thank the Lyman Entomological Museum, for providing me the
space and environment to work for my research as well as most o f the material for my
study. Moreover 1 would like to extend my gratitude to the Academy of Natural Sciences
of Philadelphia and i is cüllection manager Donald Azurna, for the loans of specimens and
the excellent cooperation we had these 2 years. 1 am furthemore grateful to the
Onhopterists' Society, for providing me with hnds for my research. McGill University
also provided a considerable amount of money through a bursary and several fee waivers;
therefore 1 would like to express rny appreciation to the Department of Natural Resource
Sciences.
Overall, 1 would Iike to thank my research supervisor Dr. CC. Hsiung, to whom 1
will dways be gratefbl. Without his help and support this work would have been
impossible to be completed. 1 want to thank him not only because of his scientific
directions, and his cooperation as a supervisor, but also for his psychologicd support and
advises in several personal problems I faced. Additionally, 1 would like to thank Dr.
Vickery for his contribution to my work. No matter day or hour Dr. Vickery was always
willing to answer my numerous questions, to help me out and to direct me the right way.
His guidance duririg my studies was extremely valuable, and 1 feel fortunate to have been
able to work with one of the world's leading authorities in Orthoptera. Moreover, 1
should also thank Dr. Wheeler, for his encouragement at my first conference presentation
in Edmonton, his endless comrnents and corrections to rny written texts, as well as his
personal cornments and suggestions about my work, which they al1 helped me become a
better systematist.
A p m for the professors directly associated with my work, 1 would first like to
thank Prof. Lockwood of University of Wyoring, for his personal support from the first
moment of our communication. ieff was the perfect colleague, who would dedicate his
time answering my melanopline-related questions and encourage me to work harder.
Thanks also to Prof. Nick Papadopoulos of Carleton University and his wife Irinela for
providing their endless support and valuable assistance whenever 1 needed.
In addition, 1 ought to thank Patrice Bouchard, Stephanie Boucher, Scott Brooks,
Paul Cermak, TilIy Gaoh, Corey Keeler, Cyrena Riley, and Rachel Wallage for being both
great colleagues and friends. Thanks aiso to Prosanta Chakrabarty for his continuous
encouragement. Special acknowledgments also extend to Marie-France Julien for her her
assistance in the French abstract, as well as for the psychological suppon during the last
stressful period before my thesis submission. Finally, 1 would like to express my deepest
gratitude to my f a d y in Greece, for supponing me both financially and psychologically
during al1 this time 1 was away fiom home.
TABLE OF CONTENTS
.USTRACT
RESUME
DEDICATION
ACKNOWLEDGMENTS
TABLE OF CONTENTS
LIST OF F I G W S
LIST OF TABLES
Page
* - II
INTRODUCTION 1
LITERATURE REVIEW 4
Systematics of the melanoplines 4
Phylogeny 5
The interna1 male genitalia as a systematic tooI in Acrididae and
Melanoplinae
MATERIALS AND METHODS
Material examined
Dissection and drawing
Abbreviations and terminology.
RESULTS
Anatomy and fiinction of the copulatory structures
Description of phailic structures for the genera studied
DISCUSSION
Range and distribution of phailic characters in the subfamily
Melanoplinae
Phylogenetic interpretations and suggestions for reclassification in the
su b famil y
Future research
CONCLUSIONS
REFERENCES
... Vl l l
LIST OF FIGURES
Figure
1.
2.
3.
4
5.
6 .
7.
8.
9.
10.
1 1 .
12.
13.
14.
15.
16.
17.
18.
19.
Intemal male genitalia of Podisma pedesrris
Intemal male genitdia of A rgiacris rehni
I n t e d male genitalia of Asemopfus montanus
Intemal male genitaiia of Bradjnotes o. obesu
Internal male genitaiia of Buckeflacris c. chifcofinae
Internal male genitaiia of Barytetfix h. hymphreysii
Internai male genitalin of Conalcaea miguelinata
Intemal male genitaiia of Dac~lotum bicolor picium
Intemal male genitalia of Hesperofettix virirüs praiensis
Internal male genitaiia of Dichropftis elongatus
Internai male genitalia of Aeoplides t. tumbulli
Int emal male genitalia of Aptenopedes sphenarioides
Internal male genitaiia of Hypochfora albû
Internai male genitaiia of Melanopius femzrrmbrz~m
Internal male genitaiia of Paroxya clavuliger
Internal male genitalia of Phoetaliotes nebrascer~sis
Internai male genitalia of Appafachia urcana
Internal male genitaiia of BohemanellaJ fi.isda
Internal male genitalia of Booneacris g. gfucialis
ix
Page
67
69
71
73
75
77
79
8 1
83
85
87
89
9 1
93
95
97
99
101
103
Figure
20 Intemal male genitalia of Dendroretrix queras
2 1 Internal male genitalia of Primnoa primnoa
22 Internal male genitalia of Pmmnacris rainierensis
Page
105
1 O7
1 O9
LIST OF TABLES
Table Page
1. Classification of the subfamily Melanoplinae, as proposed by Rehn and
Randel1 ( 1 963). I I l
2. List of genera and exemplar species studied in the present work. 113
3. Tenninoiogy used by different authors for the interna1 male genitalia of
Melanoplinae. 115
4. Comparative table of the intemal male genitalia of the melanopline genera
studied. Real proportions are rnaintained for convenience in cornparison. 1 18
Since the begiming of this century, male genitalia have been widely used as a
useful tool in systematics in most insect orders (Eberhard 1985). However in
grasshoppers, it was not until the middle 1930's that systematists included the copulatory
structures in their studies. Especially after Dirsh (1956) who proposed a new higher
classification in Acridoidea based on the phallic cornplex, more and more orthopterists
adopted the inclusion of such structures in taxonomie papers, and today this is considered
a standard procedure for most saltatorian families.
The Melanoplinae ("non striduiating grasshoppers") çonstitute the largest and
most abundant subfarnily of the New World Acrididae (Vickery & Kevan, 1986), with
more than 150 genera and a large (but undeterminecl) number of species. The immense
variability in male genitalia in the Melanoplinae has been observed by most systematists
that have dealt with members of this subfamily. This variability is believed to be due to
the fact that these structures may serve as the primary mechanism of reproductive
isolation between different species (Otte 1970, Cantrall and Cohn 1972, Cohn and
Cantrall 1 974). This is relative1 y rare arnong Orthoptera sensu stricto, where stridulation
and species-specific precopulatory behavior are usualIy the barriers to insemination
between different species. Consequently, it is likeiy that the copulatory structures of the
melanoplines could provide useful information on species limits, phylogeny and the
rnechanisms of reproductive isolation in these species.
Despite the significance of the copulatory structures, the available literature is
rather sparse and incornplete. For South American melanopline genera of Dichroplini, a
major work has been carried out over the last 3 decades (for example those of
Amedegnato 1 974, 1 977, Amedegnato and Descarnps 1 979, Ronderos and Cigliano 1 990,
199 1). On the other hand, for genera that occur north of Mexico, the overall work is
undoubtedly incomplete, since it does not include al1 genera, detailed descriptions are
ornitteci, while highly important structures are neglected. Moreover, no cornparison
between the different genera has ever been conducted. An attempt was made by Rehn and
Randell ( 1963) who illustrated some genitalic parts of several melanopline genera,
nevertheless they based their classification primarily on extemal characters, admitting that
their study was only "a tentative one".
Furthemore, the tribal classification of the subfamily lacks adequate
confirmation. Most of the characters presented by Rehn and Randell (1963) [and that
were subsequentl y followed by Otte ( 1 995) and Vickery ( 1 997) ] are overlapping, while
cytological data presented by Fontana and Vickery (1976) and Vickery (1977) include
very few genera, basically podismines. No detailed cornparison (apart fiom that of Rehn
and Randell, 1963) has ever been conducted between the tribes based on the morphology
of the concealed genitalia, even though genitalia are still - as stated above - the basic tool
used in most systematic studies concerning tribes of Acridoidea.
From al1 the above, it is now obvious that more comprehensive rnorphological
work is needed, which would give rise to usefùl information conceming the concealed
genital structures of the Nonh American non-stridulating grasshoppers.
2
Therefore, the objectives o f this project are to:
i. Describe in detail the morphology o f the interna1 male genitalia of selected North
American genera o f Melanoplinae
ii. Survey and describe the range of morphology, while defining taxonomically usefùl
differences in several characters
iii. Identifi new characters that may be useful in analyzing relationships between the
melanoplines
2. LITERATURE REVIEW
2.1. Systematics of the meianoplines
The subfamily Melanoplinae belongs to the farnily Acrididae, as do al! the true
grasshoppers (Vickery and Kevan 1983, 1986). The first classification of the
melanoplines was that of Scudder (1897a,b). He studied the genera of the "group"
"Melanopli" (spur-throated grasshoppers) of the farnily Acrididae. Later, and until the
early 1970's the narne Cyrtacanthacridinae was widely used among American authors to
include al1 acridid grasshoppers that possessed a characteristic prostemal spine. At the
same time, European authors treated sirnilar Palearctic species as sub famil y Cantatopinae.
Cantrall (1 968), Amedegnato (1974) and Cohn and Cantrall (1974) recognized a
new independent subfamily Melanoplinae, while the Cyrtacanthamidinae as recognized
today includes a smaller number of genera, known as the "bird locusts". Although the
melanoplines were thereafter widely considered as a single subfarnily (with tnbes
Melanoplini and Podismini), Otte (1995) divided this group into two different
subfamilies: Melanoplinae (Tribes Conalcaeini, Dichroplini, Dactylotini, Melanoplini)
and Podisminae.
Looking at a lower (tribal and generic) level, the complexity of the different
proposed classifications
subfamily into 3 tribes;
Randell 's ( 1 963) work,
increases. Rehn and Randell ( 1 963) suggested separation of the
Melanoplini, Podismini and
the characteristics that were
4
Dactylotini. (Table 1)- In Rehn and
used to differentiate the tribes were
so overlapping that several genera remained atypical, making the classification
" ... admittedly, a tentative one, ..." (Rehn and Randell 1963, p. 6). For two genera
(Conalcaea Scudder and Batytettix Scudder) that did not fit the existing classification,
Cohn and Cantrail (1974) erected a new tribe, the Conalcaeini. Additionally, they noted
that many genera have not been assignai to any tribe, confirming once more the generk
heterogeneity of the Melanoplinae.
In the present work, 1 am accepting the tatest published classification (Vickery
1997), who divided the subfarnily Melanoplinae into 7 tribes (Melanoplini, Dactylotini,
Conalcaeini, Bradynotini, Primnoini, Dichroplini, Podismini), basically following Rehn
and Randell's (1 963) work and elevating some groups to tribes.
2.2 Phylogeny
The phylogeny of subfamilies of Acrididae is based almost exclusively on
evolutionaq taxonomy. Most of the hypotheses are based on overall similarity of extemal
and genitalic characters combined with behavioral and ecological data. To date, there
have been only 3 studies (Cigliano 1989, Ronderos and Cigliano 199 1, Cigliano 1997)
that include cladistic methods applied to species of Melanoplinae , al1 of them involving
only a srnail number of South American genera (1 5 - 6 and 7 respectively). Consequently,
the hypotheses presented until today are al1 based on evolutionary taxonomy. This does
not necessaril y
species; indeed
imply that they might not reflect the real phylogenetic histoq of the
as Flook and Roweil(1997) state for some caeliferan superfamilies, most
5
of the morphology-based hypotheses are generally in congruence with those obtained
from curent molecular techniques and cladistic analyses.
It is generally agreed that the split between the tribes Melanoplini and Podismini
is very ancient, estimated to have occurred approximately 200 million years ago (Vickery
1987 ). The process of evolution in that group continue. before and afler the separation of
North Arnerica and Eurasia. Then, the Podismini were produced from one or more
Eurasian centers, while in the Amencas, the Melanoplini probably onginated in the
southwestern part of the North American continent. In North Amerka, members of
Melanoplini continued to speciate and disperse, producing the genus Melanoplus Stal and
its allied genera, while in South America, the genus Dichroplus Stal was produced h m
other centers of radiation (Vickery 1989). It is believed that these two genera are "sister-
genera", wry close1 y related, Melanoplus being the largest Nearctic genus and Dichrophs
being the largest Neotropical genus.
After the separation of Ewasia and North Arnerica, the Melanoplini and Podismini
started developing in isolation. The Podisrnini continued to radiate in montane regions of
eastern Asia (Rehn 1954). Fontana and Vickery (1976) suggested that, in a part of the
ancestral populations, a loss (probably by fusion) of a pair of autosomal chromosomes
divided the tribe into two subtribes Podismina Rehn and Randell ( 1 963) (23
chromosomes) and Miramellina (as group Miramellae) Rehn and Randell (1963) (21
chromosomes), both occurring widely in Eurasia. Rehn and Randell ( 1963) believed that
another group (Bradynotae, now Tribe Bradynotini) was also radiated fiom the Podismini,
including al1 podismine genera that occur in western North America. Fontana and
Vickery did not examine these genera cytologically, so their taxonomic status might
eventually be reassigned.
There has been little research to date on phylogenetic relationships between the
genera of the subfamily Melanoplinae. Al1 the papers that deal with the evolution and
classification of this group are based on similarities between genera or species to indicate
probable evol utionary pathways.
2.3 The interna1 male genitaüa as a systematic tool in Acrididae and Melanoplinae
During the late 1890's until the early 1920's, classification in Orthoptera was
based only on extemal characters such as the pronotum, prosterna1 process, furFulae,
cerci, and subgenital plates. The first complete systernatic work in the subfamily
Melanoplinae was camied out by Scudder (1 897a), who used external genitalia (together
with other external characters) to describe al1 the North American genera of the
subfamily. Two decades later, Blatchiey (1920) and Hebard (1 9 17, 1 9 18, 19 19, 1920,
1 922) presented new melanopline genera stili using extemal genitalia.
Nevertheless, Chopard (19 18) had already highlighted the value of the copulatory
structures as a taxonomic character in orthopteroids. In his paper, he provided the first
differences in the intemal genitalia of cockroaches, mantids, crickets, walking sticks and
locusts, pointing out their potential use in the classification of these groups. Later,
Chopard (1920) published his results concerning the copulatory structures of several
families of Orthoptera sensu loto, with illustrations and character descriptions, along with
his proposed terminology of the anatomy involved.
Walker (1919, 1922) followed Chopard's (1920) initial research and presented
new information concernirg male genitalia and their possible contribution in establishing
phylogenetic relationships in orthopteroids. Some new terminology was added, as well as
more species descriptions 60m a greater range of families and genera.
Hubbell (1932) gave the first remarks on the taxonomic value of the concealed
male genitalia of Melanoplinae. He noted that the vast majority of the species belonging
to the genus Melanoplus have a distinc tivel y specialized penis. Furihermore, he suggested
that the dorsal and ventral lobes of the aedeagus and the ectophallus were by far the most
taxonomically important structures. Atter Hubbeil's work, more systematists started to
look into intemal male genitalia in the melanoplines. The most striking exarnpie is
probably that of Hebard who started the first parts (Hebard 19 1 7, 1 9 1 8, 19 19, 1920, 1922)
of a revision of Melanoplinae (as "Meianopli") of United States and Canada using solely
external characters (cerci, supra-anal plates), while in al1 later parts ( 1935, 1936, 1937) he
included illustrations of genitalia in his descriptions of his work.
Meanwhile, Snodgrass ( 1 93 5) provided descriptions of the male grasshopper's
abdominal mechanisms, with detailed reference to the anatomy involved, improving and
enlarging the teminology previousl y proposed by Chopard ( 1 9 1 8) and Walker ( 1922).
His illustrations of genitalia were excellent and it is unfortunate that his study included
only one genus (Melanoplus) of Melanoplinae. However, despite the paucity of taxa
examined, he stated that "the phallic structures will be found to be more variable and
more specialized among the Melanoplini than in any other acridid group" (Snodgrass
1 935: p. 86), a fact that was thereafter confinneci by every taxonomist that studied male
genitalia in Amidoidea.
A more detailed analysis of the concealed male genitalia of grasshoppers was
presented by Roberts (1941), who conducted a comparative study of the subfamilies of
the Acrididae on the basis of their phailic parts. This was the first time that different
subfamilies were compared using male genitalia, structures of "outstanding value as
evidence for not only the diagnosis of the subfarnilies but also for their relationships"
(Roberts 1941, p. 234). Roberts (1941) included simple descriptions of genitalia, a table
of structural di fferences of the subfamilies, new addi tional terminology of male genitalia,
erection of a new subfamily (Romaleinae) and finally, a brief discussion of phylogeny in
Acrididae. He described only a single species in each of two melanopline genera
(Melanopius and Podisma Berthold), since he was including only a few (sometimes only
one) exernplar taxa for each group of aaidids.
While many taxonomists were studying the male copulatory organs, Slifer (1 939,
1 940a,b, 1 943a,b) studied the reproductive system of fernale grasshoppers. She illustrated
the female intemal genitalia (basically the spermathecae) of a wide range of species in
several subfamilies of Acrididae, including most of the North American melanopline
genera. Her results indicated the immense variability in shapes between the subfamilies
but also between the studied genera. Studies on female genitalia in Orthoptera sensu
stricto were abandoned thereafk [with the exception of Dirsh (1957), Randell ( 1963), as
well as a senes of papas in Pyrgomorphidae by Kevan et al. (1 969- 1976)], since the male
stnictures were proven to be more informative, valuable and by far more çonvenient to
study.
Dirsh ( 1956) classified the grasshopper subfamilies on the basis of their phallic
cornplex, providing extensive diagnostic keys for each group and illustrating male
genital ia of more than 700 amidoid species. In his extensive study, he illustrated only the
epiphalli of several Nearctic and Palearctic melanopline species. without providing any
descriptions. Two decades iater he presented another classification of Acridoidea,
basically by reexamining his 1956 data (Dinh 1975). His phylogenetic interpretations
were not accepted by most orthopterists (Vickery, personal communication), however the
vast majority of orthopterists use his terminology, methods of dissection and illustration.
Dinh (1 956) provided the incentive for other taxonomists to conduct further
detailed study on the genitalia of al1 acridoid families (e.g. Eades 1961, Kevan et al.
1969- 1976, Hollis 1970, 197 1, 1975, Lee 1980). Currentiy, al1 revisions of acridoid
groups that are published in the Journal of Orrhoptera Research include descriptions of
male genitalia, a fact that indicates their importance and taxonomie value (Naskrecki
1992, Roberts and Carbonell 1992, Storozhenko and Otte 1994, Jago 1994a,b, 1996,
Carbonell 1996, Gmnshaw 1996, Cigliano 1997).
After Dirsh's (1956) work the study of melanopline interna1 genitalia started to
flounsh. Gumey and Brooks (1959) described the male genitalia of the Mexicanus group
of Melanoplus, and VanHorn (1965) described the phallic morphology among
populations of Melanoplus dodgei (Thomas) in the Colorado Front Range. These two
works were undoubtedly significant fiom an ecological and taxonomical point of view,
but they were limited, covering only some species of the genus Melanoplus.
Rehn and Randell (1963) carried out ûn extensive (in terms of number of taxa)
analysis of the subfarnily Melanoplinae. Using phenetic similarities of male and f m a l e
genitalia fiom both Palearctic and Nearctic taxa, they attempted to classi* melanoplines
in several tribes and subtribes. Even though they illustrated genitalic parts (epiphalli and
some aedeagi for the males, subgenital plates and spermathecae for the fernales), their
classification was based primarily on extemal characters; unfortunately descriptions of
shared genitalic characters that would define each group were poor. Moreover, Cohn and
Cantrall (1974) highlighted the fact that "these authors, in their generic assignments,
undoubtedly depended heavily upon character similarities, as well as subjective
evaluatior, (...) of a taxon" (1974, p. 18). This fact was also emphasized by Fontana and
Vickery ( 1 976). Even Rehn and Randell (1963) admitted that their classification was
admittedly "tentative".
Despite the drawbacks, Rehn and Randell's (1963) classification is accepted even
today by some systematists. Indeed, the two latest classifications of Melanoplinae
recognized the tribes Dactylotini, Dichroplini and Melanoplini (Otte 1995) as well as
tribes Bradynotini and Pnmnoini (Vickery 1997) which were first suggested by Rehn and
Randell (1 963) as "groups" (meaning subtribes) of the tribes Melanoplini and Podismini
respectively.
From the late 1960's through the 703, the taxonomie effort in Melanoplinae was
significantly stronger, in terms of quantity and quality of papers published. Genitalia were
(and still are) the most effkctive tool in melanopline systematics due to their variability.
Ronderos and Carbonell worked mostly with the Dichroplini, a South American group of
melanoplines and published a great number of papers, al1 based on intemal male genitalia
(Ronderos 1964, Ronderos et al. 1968, Carbonell and Ronderos 1973, Rowell and
Carbone11 1977, Ronderos 1979a,b, Ronderos 198 1, Ronderos and Cedra 1982). Many
species in several genera of Dichroplini can only be identified by differences in genitalia,
so their work revealed numerous new species and subspecies. Similar detailed studies for
the North Amencan melanoplines on the bais of their phallic structures has not been
canied out since Rehn and Randell(1963).
Amedegnato, who also worked primaril y with South American melanopline
genera, published a series of papers (Amedegnato 1974, Amedegnato and Descamps
1978, 1979), in which classification and phylogenetic relationships were established
basically upon male genitalic characters. She was the first to recognize the independence
of the subfamily Melanoplinae (Amedegnato 1974). The subfamily was based on the
presence of a constricted point of articulation between the apical and basal parts of the
aedeagus, cornpared with the presence of a sigmoid flexure in the Catantopinae. This was
previousl y observed by Roberts ( 1 94 1 ) for the genera Melanoplus and Catuntops Schaurn
respectively, and was later confirmed and accepted for the Melanoplinae by Cohn and
Cantrall ( 1974).
Cohn and Cantrall (1974) suggested that the differences occumng in the male
genitalia of the melanoplines were
of the species involved. They took
exclusive1 y responsible for the reproductive isolation
into account the fact that in Melanoplinae, acoustical
12
and visual precopulatory behaviors are not highly developed, in contrast with other
subfarnilies of Acrididae. As Otte (1970) has stated, fernales in Cyrtacanthacrinae
(=Melanoplinae) are taken "by surprise" by males and the males may be rejected after
mounting (if they do not beiong to the same species) or not, while in other subfarnilies
preîopulatoiy behaviors occur before mounting so that recognition of the sarne species is
achieved and thus copulation is t!!ereafier without rejection of males.
Based on this fact, Cohn and Cantrall (1974) confirmeû that the copulatory
structures in Melanoplinae are under strong selection, holding particularly valuable clues
about the biological and phylogenetic interrelation of the species involved. More
specifically, Cohn and Cantrall(1974) proved that in the genus Baytettix, the only barrier
to insemination between différent species is the misfitting of the genitalia and thus there
is a direct selective action on the genitalia as a reproductive isolation mechanism.
Vickery and Kevan (1983, 1986) described al1 the Canadian genera of
Melanoplinae, but genitalic descriptions were omitted. This was probably due to the fact
that external genitalic parts, such as the subgenital plate, the cerci, and supra-anal plate,
were (and still are) good characters for species identification and are easier to use by a
non-taxonomist. However, Vickery and Kevan (1983, 1986) noted that the phallic
complex constitutes the only reliable way to separate some species in which external
characters overlap with each other (e.g. Melanoplus huroni Blatchley and M. dodgei
(Thomas), M. foedus foedus Scudder and M. packardii Scudder). In fact it is probable that
the examination of phallic structures
increase the number of species and
in North Amencan Melanoplinae will definitely
subspecies in the subfamily (Vickery, personal
13
communication).
Whether or not genitaiic difierences should be the basis for erecting a new species
was a main point of argument between Lockwood (1996) and Cohn (1994), a discussion
which rejuvenated the interat in studying melanopline genitalia in the 90's. Lockwood
(1996) clearly indicated that reliance on the aedeagus has little validity because of its
possible intraspecific variation, as well as its correlation with ecological conditions. Cohn
(1994) additionaily stated that genitaiia (despite his strong support about their usehlness
and informative power) are insufficient to define species, since "...most times they are
good species discriminators, sometimes they are not" (Cohn 1994, p. 59). However he
States: "1 am unaware of any orthopterist who explicitly rejected the use of genitalia in the
Melanoplinae" (Cohn 1994, p. 60). Overall, it should be realized that any aection of a
new species based on morphological characters (extemal or genitalic) consists of a
hypothesis. that thereafter should be confirmed by additional morphological, molecular,
behavioral and biogeographical data.
3. MATERIALS AND METHODS
3.1 Material examined
In the present study, exemplar taxa were selected to represent each group. The
selection of genera was based on: i) their geographical distribution (to inctude al1
Canadian genera), ii) their classification (to include all tnbes and subtribes), iii) the
availability of specimens mainly at the Lyman Entomological Museum and Research
Laboratory and Academy of Natural Sciences of Philadelphia. Obviously, the choice of
genera could have been more extensive, nevertheless 1 hope the results fiom this work
will be informative enough to draw the basic lines in the subfamily. The selection of
species for each genus was mainly based on the type species, as well as their availability.
However, it was realized that a study of more than a single species for each genus would
probably provide more usehl information. Apart fiom the genera Aeoplides, Baryiettk,
Dichroplus, Appalachia, the rest of the 18 genera are represented by their type species
(Table 2).
Most specimens for this study were obtained from the Lyman Entomological
Museum and Research Laboratory. Specimens that were obtained for the Academy of
Natural Sciences of Philadelphia are indicated by "ANSP".
Argiacris rehni Hebard
1 ) USA, ID1 Galena Summit, Sawtooth Range. 8750 to 9225 feet. 17-ViII-1928. (R & H)
(3 males, ANSP); 2) USA, MT, Livingston, 03-VII- 190 1. (1 male, ANSP)
Asemoplus montanus Bruner
1) USA, ID, Lemhi Pass, Lemhi Co. 1 1 -VI[- 1956 (W.F. Barr) (3 male paratypes, ANSP);
2) CANADA, AB, Swan Range, Mont., 2900 feet.. 14-VIII-1905 (M.J. Elrod) (1 male
paratype, ANSP); 3) CANADA, BC, Rock Creek, 25-VIL 192 1 (Buckell) (1 male, ANSP)
Bradynotes obesa obese Thomas
USA. MT, Heeler, (3 males, ANSP)
Buckellacris chilcotinae chilcotinae Hebard
1) CANADA, BC, Old Green Min, Ofd Apex Mtn Rd., 18-VI-1982, Kirk, (2 males); 2)
CANADA, same locdity and collecter as in ( 1), 25-VI- 1982, ( 1 male); 3) CANADA, BC,
Apex Mtn, near Keremeos, 26-VI-1982, (1 male); 4) CANADA, BC, Douglas Lake, 27-
VI14924 (E.R. Buckell) (1 male); 5) CANADA, BC, Kelowna, 18-VIL1927 (E.R.
Buckell) ( I male)
Baryte& hymphreysii hymphreysii Thomas
1) USA, AZ, , Atascosa Mountain, Pajaritos Mts., Santa Cruz Co., 5,000-6,500 feet, West
foot of peak, 214x4922 (H) (2 males, ANSP; 2) USA, AZ, Sonora Road Canyon,
16
Tucson Mts, Pima Co, 3,000 feet, 1 1 -X- 19 10 (1 male, ANSP); 3) USA, Sabino Basin, St.
Catalina Mts., 32722' N. 1 10°46,5' W, about 3,800 feet, 8,20-VII-1916 (1 male, ANSP);
4) USA, AZ, Sycamore Cadon, Baboquivari Mts., Pima Co., 4,700 feet, 6,9-X-19 10 (1
male, ANSP); 5) USA, AZ, Phoenix, (R.E. Kunze) (1 male, ANSP); 6) USA, Bear
Valley, Pajaritos Mts., Santa Cruz Co., 5,500 to 5,700 feet, 21-IX-1922 (R.) (1 male
ANSP)
Conalcaea miguelinata Scudder
1) MEXICO, between Zacapu and Zamora, Mich., 7500 feet. 6 4 x 4 9 3 8 (H.R. Roberts)
(7 males, ANSP); 2) MEXICO, 5 miles south of Chilchota, rd. to Uruapan, Mich., 6000-
7000 feet, 15-VIII-1940 (H.R. Roberts) (2 males, ANSP); 3) MEXICO, 18 miles West of
San LuisPotosi, S.L.P., 7500 feet, 5-K-1940 (1 male, ANSP)
Ductylotum bicolor pictum Thomas
1) USA, NE, MomlI Co. 2-VIII-1958 (R.H. & E.M. Painter) (1 male) 2) USA, CO,
Brash Hollow, 20-1x4970, (G. Bauzner) (1 male); 3) USA, NM, Clifion House, Colfax
Co., 6360-6400 feet, 4-VIII-1921 (R) (1 male, ANSP); 4) USA, NM, Clifton House,
Colfax Co., 6355 feet, 12-VIL1937 (Rehn, Pate & Rehn) (2 males, ANSP); 5) USA, KS,
Grant Co., 2800 feet. 23-VIi-19 1 1 (F.X. Williams) (2 males, ANSP)
Hesperotettir viridis pratensis Scudder
CANADA, AB, 10 mi. E Fort Macleod, 9-VIII- 1967 (Colin & W. Boyle) (7 males)
17
Dich roplus efongatus Gi g 1 io-To s
1) ARGENTXNA, San Juan, Prov. San Juan, 673m, 17-1-1909 (P. Jorgensen) (1 male,
ASNP; 2) ARGENTiNA, Jujuy, Cornell University Expedit., 10-11- 1920 (R.G. Harris) (1
male, ANSP); 3) ARGENTINA, Maria Lulila F.C.M., 18-1-1939 (C.H. Richardson) (2
males, ANSP); 4) ARGENTINA, Tigre, Buenos Aires, Cornell University Expedit., 8-11-
1920 (2 males, ANSP); 5) CHILE, El Olivar Colchagua. (C.S. Reed) (1 male, ANSP); 6)
CHLE, Laugostas Las Caudes, 3-VI-1965 (2 males, ANSP); 7) South America, Tucwnan,
Est. Expt. Agric., 24-IV- 19 13 (2 males, ANSP)
Aeopiides turnbuiü turnbuUi Thomas
1) USA, MT Glendive, , Dawson Co, 26-ViI-1909 (1 male, ANSP); 2) USA, WY,
Worland, VI1 (2 males, ANSP)
Apten opedes sphenarioides S cudder
1) USA, FL, Gainesville, 12-X-1979 (V.R. Vickery) (1 male) 2) USA, same locality and
collector as in ( 1 ), 14-X- 1979 (2 males); 3) USA, same locality and collector as in (1 ), 10-
X- 1979 (1 male); 4) USA, same locality and collector as in ( 1 ), 5-XII- 1979 (2 males); 5)
USA, FL, Welaka Reserve, 90x14979 (V.R. Vickery) (2 males); 6) USA, FL, Broword
Co., Hallandale Beach, 24-XI-1961 (D.K.McE. Kevan) (1 male); 7) USA, FL, Broword
Co., Everglades Pk. Palma Visha Hammock, 26-XI- 196 1 (D.K.McE. Kevan) (1 male)
Hypochlora al& (Dodge)
1) USA, Western USA, 244x4973 (K. Hofban) (1 male); 2) USA, Western USA, 18-
X-1973 (K. Hofhan) (1 mate) 3) USA, MT, Colombus, 1 -1X-1924 (N. Criddle) (1
male)
Melanoplus fernmurrubrum (De Geer)
1 ) CANADA, QC, Ste-Anne-de-Bellevue, 24-VIII-1967 (W. Boyle) (2 males); 2)
CANADA, same locality and collector as (l), 26-VI114967 (7 males); 3) CANADA,
same locality and collector as (l), 84x4967 (1 male); 4) CANADA, same locality as (l),
5-IX- 1967 (R. Lalonde) (3 males); 5) CANADA, BC, Salmon Ann, 1 8-VIII- 1966 (V.R.
& W.L. Vickery) (1 male); 6) CANADA, SK, Blackstrap Valley, 3 min. Dundurn, 23-
VIII- 1 966 (V.R. Vickery) (1 male); 7) CANADA, ON, Hawkesbury, 2 1 -1X- 1963 (V.R.
Vickery) (1 male); 8) CANADA, AB, Lethbridge, 28-VII-1967 (C. & W. Boyle) (1
male); 9) CANADA. MB, Lyleton, 12-VIII-1967 (C. & W. Boyle) (1 male); 10)
CANADA, NS Collingwood, Cumb. Co., 15-IX-1959, (V.R. Vickery) (1 male); 11) USA,
COI Grand Junction, 12-VIL 1966, (W. Boyle & R. Lalonde) (1 male); 12) USA, NC,
Rest area, 1-77, Va-N.C. border, 224x0 1979 (V.R. Vickery) (1 male)
Paroxya clavuliger ( Audinet-Servi lle)
1) USA, FL, Monroe Co. Pinecrest, Route 94, 244-1968 (Boyle, Lalonde & Ng.) (4
males); 2) USA, FL, Dade Co. Jct. 27 and 94, 17-V-1968 (Boyle, Lalonde & Ng.) (1
male)
Phoetaliotes nebrascencis (Thomas)
CANADA, AB, 10 mi. E Fort Macleod, 9-VIII- 1967 (Colin & W. Boyle) (1 O males)
Appalachia arcana Hubbell and Cantrall
1) USA, MI, Oscoda Co., Sec.20, 6 min. NW of Mio, June. Co. 608 & Yomgs Rd., 13-
VIII-1974 (Crozier, Fontana & Ladd.) (5 males); 2) USA, MI, Oscoda (o., Sec.2, 5.5mi.E.
on Co489 off M33 (8 min. SE of Mio), 12-VIII- 1974 (Crozier, Fontana & Ladd.) ( 1 male)
Bohemaneila frigr'da fn'gtia Boheman
1 ) CANADA, YT, Canoe Lake, 68'13' N.:l35"5S7 W, 10-VIII-1965 (D. Momson) (4
males) 2) CANADA, NT, Eskimo Lakes, Mackenzie, 24-VIII- 197 1 (J. Lovrity) (1 male);
3) FRANCE, Col du Grandon (H. Alpes), 24OOM., VIII- 1 965 (G-Remaudiere) (2 males)
Booneacrk glacialis gkrcialis Scudder
CANADA, QC, Mt. Albert, Gaspesian Park, 15-VIII- 197 1 (Boyle & Ladd) (4 males)
Dendrotettix quercus Packard
U S A , MI, Lake Co., 4 min. N of Baldwin on M37, 13,14-VIII-1974 (Crozier, Fontana &
Ladd) ( 1 2 males)
Podisma pedestrls Limaeus
1 ) GERMANY, Bavaria. Berchtesgaden NP., 950 m., 30-VIII- 1984 (V.R. Vickery) (6
males) 2) FRANCE, entre Sault & le Chalet Reynard (Vaucluse) 1300 M., 22-VIII-1967
(M-Descarnps) (1 male); 3) FRANCE, Mt. Serein (Vaucluse) 1500 M., 4-VIII-1967
(M-Descarnps) ( 1 male)
Primnoa primnoa Fischer-Waldhein
1 ) RUSSIA, Postweg Amginsk - Jakrtsk, 14-V1II-1925 ( 1 male, ANSP); 2) RUSSIA,
Amur. Oblast, ca.150km north of Blagoveshchensk, Tukunngra Mts, 8-1x4956 (D.
Koronov) (1 male, ANSP); 3) RUSSIA, 5-VIII-1925 (1 male, ANSP); 4) RUSSIA, Stat.
Hingran Mardzhavria, 5411-1925 (1 male, ANSP); 5) RUSSIA, 44'30' N., 130' E,
Greenw., 22-VI- 1 9 1 0, (N. Ikonikov) ( 1 male, ANSP)
PrumnacriS rainierensis Caudell
1 ) USA, WA, Mount Rainier, Pierce Co., Camp o f the clouds, 5400-7200 feet, 23,24-
VIII- 19 10 (8 males, ANSP); 2) USA, WA, Mount Rainier, Pierce Co., Lower Paradise
Val. Above Narada Falls, 4500 feet, 23,24411- 19 10 (1 male, ANSP)
3.2 Dissection and drawing
For dissections, the techniques of Roberts ( 194 1 ) were followed, with
modifications by later workers (Dirsh 1956, Rehn and Randell 1963).
Relaxation. The specimens fiom the museum collections were dry, so their abdomens had
to be relaxed pnor to dissection. Dirsh's (1956) methods, which requires the specimens to
be left for 24 hours in a relaxing chamber, containing a few drops of ethyl acetate diluted
in water, was initially used in the present work, but was abandoned since it caused
discoloration of the specimen. Instead, the distal part of the abdomen was submergeci 4 to
5 hours in warm water.
Positioning. After relaxation, the specimen was positioned in a wax-bottom dish filled
with water and placed under a binocular microscope for dissection. Wax provided a firm
surface for the specimen to be pinned and water allowed easy manipulation of the phallic
part and membranes. The specirnen was placed with the ventral side down and several
pins were used to spread the wings and legs on each side.
Dissecting. In this stable position, a needle was used to depress the subgenital plate.
Subsequently, the pallium was cut Iaterally and pulled back to uncover the phallic
complex. The phallus together with the epiphallus were rernoved with a pair of fine
forceps and after cutting the membrane surrounding the base of the phallus, the whole
phallic complex was extracted. This procedure ensured that the extemal parts (subgenital
plate, cerci, supra-anal plate) remained undmaged. These extemal genital structures had
to be returned to their normal positions, as they are essential for species identification and
therefore should appear unaftered.
Muscle rernoval. The phallic complex, together with its surrounding membranes were
placed into a small g l a s via1 containing a 5% solution of KOH. The via1 was placed on a
hot plate at approximately 150" C for about 10 min, time enough for al1 muscles to be
dissolved. The çolor of the remaining sclerotized parts was sometimes darkened, but this
did not affect the shape of the structures, nor did it obstruct the drawing procedure.
Dissection of sclerotized parts. First, the membrane comecting the epiphallus to the rest
of the phallic complex was cut, so that these parts could be drawn separately. The minute
membrane between the cingulum and the aedeagus also had to be cut, around the arch of
cingulum, in order to separate these two structures.
Drawing. Drawings were made with the use of a carnera lucida on a WILD M5 dissecting
microscope. Each phallic part was pIaced into a drop of glycerol on a slide, and
positioned with a pair of needles. The glycerol prevented the movement of the structure,
so that it remained stable for fûrther drawing.
Al1 figures follow the semidiagrammatic pattern used by Dirsh (1956) and other
authors (e-g. Roberts 194 1, Rehn and Randell 1963, Kevan et al. 1969- 1976), in which
details such as superficial scuipting are omitted. Only the sclerotized parts and the most
important membranes are illustrated, disregarding parts such as the ejaculatory duct, the
ejaculatory sac and spermatophore sac, that are not taxonomically important.
Generally, the phallic structures of a species are difficult to draw in a consistent
way, since a slight change of the angle of view may cause a different outline (Dirsh
1 956). It should therefore be realized that the illustrations presented in this work might
differ from the actuaI preparations or fiom figures of other works as they might be viewed
fiom different angles. However, the approach is vatid, since it is not observer biased and
could be repeatable.
3 3 Abbreviations and Terrninology
The terms used to describe the anatomy are adopted tiom Dirsh (1 W6), since they
are the ones most common!y used in acridoid systematics. Several other publications refer
to the phallic complex of Acridoidea (e-g. Chopard 1920, Walker 1922, Hubbell 1932,
Snodgrass 1935, Roberts 1941), but the change of terminology in each work makes it
difficult for the reader to foliow. Therefore, a complete list of the abbreviations that will
be used in this study together with their explanations, is presented in Table 3. The
terminology used by other authors is also indicated, in order to enable one to compare
with previous studies in melanopline genitalia. The terni "inner projections of the
zygoma" is used here for the tirst time, to describe a pair of projections extending h m
the inner side of the zygoma that "hold" the arch of cingulum.
4. RESULTS
4.1 Anatomy and function of copulatory structures
For the genera studied, the phallic complex is revealed when the pallium is pulled
back and the subgenital plate is lowered. Its apical part is directed towards the posterior
end of the body, inside the 1 s t abdominal segment. The phallic organ represents the distal
end of the ejaculatory duct and could be divided to two main parts: the aedeagus (Figure
1 B,C,E and the cingulum (Figure 1 B-E). The former is a simple widened continuation of
the ejaculatory duct which is composed of 3 parts (Figure 1E): i) the ejaculatory sac, ii)
the spermatophore sac, and iii) the phallotreme. Sperm and spermal fluid is stored in the
ejaculatory sac, whîle the spematophore that will be transmitted to the female is formed
in the spermatophore sac. These two sacs are connected by the gonopore. Final! y the male
spermatophore passes through the phallotreme and is deposited into the female genital
chamber during copulation. The two most important factors for the transmission of the
spermatophore are the movement of the skeletal part of the aedeagus (and partly the
cingulum), as well as the hemolymph pressure which is responsible for the erection and
swelling of the phallic cornplex. Other possible factors that affect the penetration of the
male genitalia into the female might be the active movement of spematodesms (Pickford
and Gillot 197 1).
The parts considered in taxonomy are the sclerotized parts of the phallic complex.
They consist of a pair of sclerites of various shapes (the apicai and basal parts of the
aedeagus) and they have several fûnctions: a) regulate and direct the spenn flow; b)
facilitate the transmission of the spermatophores; and c) penetrate the female genital
chamber. The basal parts of the aedeagus in the Melanoplinae possess a mechanism,
known as the gonopore process, that regulates the movement of the spenn fiom the
ejaculatory sac to the sperrnatophore sac (Figure I E).
The second part of the phallic organ is the cingulum, which provides mechanical
support to the phallic complex. It is a strongly sclerotized structure on the dorsal side of
the penis. It consists of a pair of elongated scterites (the apodemes) which are joined at
their proximal ends by a bar-like structure, the zygoma. The lateral processes of the
cingulum are called rami and form a sheath that envelopes the apical valves of the penis.
Finally, at the apical part of the cingulum, the valves of the cingulum are formed, usually
parallel and dorsal to the apical valves of the penis. These vaIves are joined with the
zygoma and with the arch of cingulum (Figure 1 C, E)
An organ on the dorsal side of the phallic complex is the epiphallus (Figure 1 A),
derived fiom the ectophallic membrane (membrane on the cingulum) and strongly
attached to it. It is comprised of the ancorae, the anterior projections, the bridge, the lophi
and the posterior projections. It may not play an active role in the movement of the phallic
organ, however it stabilizes the phallus and controls the positioning of the distal parts of
the penis in order to
significant structure in
penetrate the female genital chamber. The epiphallus is d s o a
taxonomy, due to the variable shape of its parts.
26
Detailed illustrations of the phallic parts of a melanopline grasshopper are
presented in Figure 1. Some structures attached to the aedeagus (such as the ejacuiatory
duct, sac and spermatophore sac, gonopore process) are not included in the rest of the
figures, since they are non-informative in an intergeneric çomparison; they are usually
used in cornparisons of families or superfamilies (Dirsh 1956, Kevan et al. 1969- 1976).
4.2 Description of phaüic structures for the genera studied
The description of the studied genera is derived fiom the illustrations presented in
Figures 1-22. The descriptions of apodemes, zygoma, and inner projections of the
cingulum were based on the fiontal view (D) and not on the dorsal view (B) that might be
confùsing for these structures. The terms "small, medium, large", or "normally
developed" that are used throughout the descriptions are relative and apply to the general
characteristics of the subfamily.
Tribe Bradynotini
A rgiacris rehni (Figure 2)
Epiphallus: Size small; ancorae medium length, with obtuse tip, slightly convergent;
anterior processes slightly longer than ancorae, acute tip; bridge two times longer than
broad; lophi round, cylindncal, relatively short, two times longer than broad; posterior
processes wide, slightly convergent, with acute tip extending laterally; imer margin of
bridge curved upwards, with small hump in the middle.
Phallic complex (except epiphallus): apodemes wide, moderately long, parallel; d e r
projections of cingulum medium size, slightly convergent; sheath of penis covering 30%
of arch of cingulum and 90% of apical valves of cingulum, extending from middle of rami
to posterior margin of distal part of rami; rami of cingulurn normally developed, with
acute distal part, Aedeam: arch of cinguium rectangular shape; apical valves of
cingulum slightly developed, rather short, reaching 50% of length of apical valves of
penis; apical valves of penis long, slender, with sigmoidal apex curved upwards.
Asemoplus monûanns (Figure 3)
Epiphallus: Size small to medium; ancorae with wide obtuse tip, convergent; anterior
processes as long as ancorae, slender, with acute tips; bridge two times longer than broad;
lophi cylindrical, elongate, at least four times longer than broad, partially protruding fiom
posterior processes; posterior processes practically non existent; imer margin of bridge
curved upwards, with no excavated or projecting parts
Phallic complex (except epiphallus): apodemes slender, relatively short, curved inwards at
distal ends; inner projections of cingulum small, angular, strongly convergent; sheath of
penis covering 90% of apical valves of cingulum and penis, and al1 arch of cingulum,
extending from zygoma to anterior margin of distal end of rami; rami of cingulum very
long, highly expanded caudally, widened at the distal part. Aedeagus: arch of cingulum
with small acute denticle at the middle of the posterior margin; apical valves of cingulum
well developed, reaching the end of apical valves of penis; apical valves of penis with a
small hump near the apex, which covers part of apical valves of cingulwn.
Bradynotes o. oksa (Figure 4)
Epiphallus: Size large; ancorae cylindrical, with round tip, convergent, well developed;
anterior processes as long as ançorae, round apex; bridge three times longer than wide;
lophi ellipsoid, two times longer than broad, strongly protniding from lateral plates;
posterior processes slightly developed, divergent round apex; imer margin of bridge
horizontal
Phallic complex (except epiphallus): apodemes relatively short, divergent; inner
projections of cingulum strongly reduced; sheath of penis covering al1 the arch, apical
valves of cingulwn, and 80% of the apical valves of penis extending fiom zygoma to
posterior margin of the distal end of the rami; rami of cingulum long, wide in lateral view,
nomally developed. Aedeagus: arch of cingulum very wide, with small acute denticle at
the top of the posterior margin; apical valves of cingulum slightly developed, very acute,
reaching 60% of the length of the apical valves of penis; apical valves of penis robust,
with wide apices, curved laterally towards rarni, serrate ventral margin.
Buckellacrk c chilcotinae (Figure 5)
Epiphallus: Size large; ancorae wedge shaped, with acute tips, strongly convergent;
anterior processes robust, as long as ancorae, v q thick and wide, with obtuse tip; bridge
narrow, three times longer than broad; lophi protruding fiom lateral plates, cylindrical,
three times longer than wide; posterior processes long, slender, with obtuse apices.
strongly divergent; inner margin of bridge slightly c w e d upwards.
Phallic complex (except epiphallus): apodemes reiatively short, divergent, flattened
apically; zygoma with long concave ridge that seems to separate the zygoma in two
lobes; inner projections of cingulum long, strongly convergent, acute; sheath of p a i s
covering approximately 70% of arch and apical valves of cingulum, and apical valves of
penis, extending fiom zygoma to anterior margin of distal end of rami; rami of cingulum
very long, slender. Aedeagus: arch of cinguium with long projection at the posterior
margin; apical valves of cingulum and apical valves of penis very complex, as in figure.
Tribe Conalcaeini
Baryte& h. hymphreysii (Figure 6)
Epiphallus: Size large; ancorae long, slender, with acute tip, strongly convergent; anterior
processes siightly shorter than ancorae, wider, obtuse tip, slightly convergent; bridge at
Ieast two times longer than broad; iophi strongiy extending fiom lateral plates,
cylindrical, parallel to inner margin of bridge; posterior processes well developed,
slender, parallel; imer margin of bridge curved upwards.
Phallic complex (except epiphallus): apodemes slender, elongate; imer projections of
cingulum well developed, rather wide; sheath of penis covering arch and apical valves of
cingulum, and 80% of apical valves of penis; rami of cingulum long, strongly developed
caudally. Aedeagus: arch of cingulum well developed, apex towards basal parts of penis;
30
apical valves of cingulum curved upwards; apical valves of p a i s slender, long, slightly
cwved upwards.
Conaîcaea migueîina~rr (Figure 7 )
Epiphallus: Size medium; ancorae long, wide, convergent, with obtuse tips; anterior
processes long, slender, acute tips, strongly convergent; bridge slightly longer than broad;
lophi slender; posterior processes long, slender, parailel, slightly convergent apices; imer
rnargin of bridge horizontal.
P hallic complex (except epi phallus) : apodemes short, with wide apices; imer projections
of cingulum very small, acute; sheath of penis mvering arch of cingulum, 90% of apical
valves of penis; rami of cingulum wide, long. Aedeagus: arch of cingulum with small
acute denticle extending h m the posterior margin; apical valves of cingulum indistinct;
apical valves of penis, long, bifùrcate, with two acute apices.
Tribe Dactylotini
Dacîylotum bicofor picrum (Figure 8)
Epiphallus: Size medium to large; ancorae small, obtuse tip, strongly convergent; anterior
processes strongly reduced, obtuse; bridge two times longer than broad; lophi long,
slender, C-shaped, with posterior apices widened, at least four times longer than broad;
posterior processes practically non existent; inner margin of bridge rather horizontal, with
excavated part in the middle, containing a small hurnp.
Phallic complex (except epiphallus): apodemes slender, long, slightly divergent, slightly
widened apices; imer projections of cingulum small, strongly convergent, acute tip,
bearing a pair of longer parallel projections; sheath of penis covering al1 apical valves and
arch of cingulum, and 50°/a of length of apical valves of penis, extending fiom zygoma to
posterior margin of distal end of rami; rami of cingulum short, nomally developed,
short, obtuse end. Aedeagus: arch of cingulurn relatively small; apical valves of cingulum
long, weli developed, acute tip, reaching 70% of the length of apical valves of penis;
apical valves of penis long, with acute tip.
Hesperotettix viridiS pratensis (Figure 9)
Epiphallus: Size medium; ancorae long and wide, slightl y convergent apical1 y; anterior
processes slender, as long as ancorae, with obtuse tip; bndge nearly 4 times longer than
broad; lophi as in Figure 9A, at least three times longer than broad; posterior processes
wide, acute tips, divergent apically; imer margin of bndge curved upwards.
P hallic complex (except epiphallus) : apodemes slender, long; zygoma short, wide; imer
projections of cingulum absent; sheath of penis absent; rami of cingulum very wide, long,
strongly extending caudally, longer than apical valves of penis. Aedeagus: arch of
cingulum srnaIl, with large acute denticle extending fiom the posterior margin; apical
valves of cingulum as long as apical valves of penis; apical valves of penis short.
Tribe Dichroplini
Dichroplus elongatus (Figure 10)
Epiphallus: Size small to medium; ancorae long, slender, finger-like, convergent; anterior
processes slightly longer than ancorae, robust, very wide, rather parallel; bndge three
times longer than wide; lophi small, oval, two times longer than wide; posterior processes
long, obtuse tip, slightly divergent; imer margin of bridge horizontal.
P hallic compiex (excçpt epiphallus) : apodemes slender, long; inner projections of
cingulum obtuse, well developed; sheath of p a i s covering 20% arch of cingulum, 80% of
apical valves of penis, extending from zygoma to anterior margin of rami; rami of
cingulum nomdly developed. Aedeagus: arch of cingulum wide; apical valves of
cingulum short, with acute apex; apical valves of penis very long, apex curved 90'
upwards.
Tribe Melanoplini
A eoplides & turnbull (Figure 1 1 )
Epiphallus: Size medium to large; ancorae wide, long, convergent, with obtuse tips;
anterior processes long, wide, as long as ancorae, wedge shaped, convergent; bridge at
least two times longer than broad; lophi small, cylindrical; posterior processes wedge
shaped, divergent with acute tips; i ~ e r margin of bndge horizontal, with minute denticle
in the middle.
Phallic complex (except epiphallus): apodemes slender,
projections of cinylum obtuse, wide; sheath of penis small,
moderately long;
covering 70% of
inner
apical
valves of cingulum, 40% of length of apical valves of penis; rami of cingulum slender,
iong; arch of cingulum with small obtuse denticle extending fiom the posterior margin.
Aedeagus: apical valves of cingulwn absent; apical valves of penis long, slender, cwed
upwards 90°, with bifhrcate apex.
Aptenopedes sphenurioides (Figure 1 2)
Epiphallus: Size small; ancorae sigrnoid shape, convergent, with acute tips; anterior
processes slightly shorter than ancorae, obtuse, divergent; bridge hook-like shape,
membrane-like texture that seems to separate the lateral plates; lophi hook form; posterior
processes very long, slender, strongl y divergent; inner margin of bridge sli ghtl y curved
upwards.
Phallic complex (except epiphallus): apodemes very long and slender, converging
apically; inner projections of cingulurn absent; sheath of p a i s covering ail arch of
cingulum, 60% of length of apical valves of cingulurn and penis; rami of cingulum
normalIy developed, very wide. Aedeagus: arch of cingulum wide, robust; apical valves
of cingulum long, slender, longer than apical valves of penis; apical valves of penis
slender, slightly reduced.
Hypochfora alba (Figure 13)
Epiphallus: Size small; ancorae angular shape, acute tip, slightly convergent; anterior
processes longer than ancorae, wide, obtuse tips; bridge two times longer than broad;
lophi ellipsoid, at ieast three times longer than broad, slightly protmding form lateral
plates; posterior processes small, wedge shaped, strongly divergent; imer margin of
bridge rather horizontal, with small hurnp in the middle.
Phallic complex (except epiphallus): apodemes relatively short, slender, slightly
divergent; imer projections of cingulurn small, obtuse; sheath of p a i s covering al1 arch
and apical valves of cingulum, 50% of apical valves of penis, extending fiom middle of
Rrn to ventral margin of distal end of rami; rami of cingulum long, widened apices in
Iateral view, slightly expanded caudally. Aedeagus: arch of cingulurn with small obtuse
denticle extending fiom the posterior margin; apical valves of cingulum reduced; apical
valves of penis, sigrnoid distal part
Melanoplus femmurubrum (Figure 14)
Epiphallus: Size large; ancorae long, well developed, convergent; anterior processes as
long as ancorae, robust with acute tip bridge two times longer than broad; lophi upcurved
with angular apices, at least two times longer than broad; posterior processes well
developed with acute tips; inner margin of bridge curved downwards, with small
excavated part in the middle.
Phallic complex (except epiphallus): apodemes very wide, long, slightly convergent; imer
projections of cingulum reduced; sheath of penis covering arch, apical valves of
cingulum, apical valves of penis and more than 60°h of length of apical valves of
cingulum, extending fiom middle of rami to posterior margin of distal end of rami; rami
of cingulum slightly short, robust. Aedeagus: arch of cingulum with large denticie
extending fkom the posterior margin; apical valves of cingulum long, widened apically,
as long as apical valves of penis; apical valves of penis long, widened apically.
Paroxya clavuliger (Figure 15)
Epiphallus: Size medium; ancorae quite long, convergent with acute tips; anterior
processes three times longer than ancorae, parallel, with acute tips; bridge two times
longer than broad; lophi lobifom, two times longer than broad, slightly protruding from
lateral plates; posterior processes slender, strongly divergent, obtuse tip; inner margin of
bridge horizontal.
Phallic complex (except epiphallus) : apodemes rather wide, short, convergent; inner
projections of cingulum absent; sheath of penis covering apical valves of cingulum, 80%
of length of apical valves of cingulum, 70% of length of apical valves of penis, extending
fiom zygoma to ventral margin of distal end of Rm; rami of cingulum wide, extending
caudally. Aedeagus: arch of cingulum small, with obtuse denticle extending fiom the
posterior margin; apical valves of cingulum very long, strongly curved downwards,
almost reaching the apex of apical valves of penis; apical valves of penis strongly curved
downwards with acute tips.
Phoetaliotes nebrascensis (Figure 1 6)
Epiphallus: Size small; ancorae long, wide, with obtuse tip convergent; anterior processes
parallel, as long as ancorae with obtuse tip; bndge very narrow, at least four times longer
than broad; lophi strongly protruding fiom lateral plates. wide, almost parallel to bridge,
two times longer than wide; posterior processes long, strongly divergent with obtuse tip;
inner margin of bridge curved upwards.
Phallic complex (except epiphallus): apodemes very long, slender, imer projections of
cingulum reduced, obtuse; sheath of p a i s covexing arch of cingulum, 50%1 o f length of
apical valves of penis, extending from zygoma to posterior rnargin of rami; rami of
cingulurn wide with obtuse distal paits. Aedeagus: arch of cingulum small, acute; apical
valves of cingulum absent; apical valves of penis extremely long, slender, c w e d
upwards.
Tribe Podismini
Appalachia orcana (Figure 1 7 )
Epiphallus: Size medium; ancorae medium length, with acute tip, strongly convergent;
anterior processes slightiy developed, shorter than ancorae, strongly obtuse; bridge two
times longer than broad; lophi lobiform with ventral margin c w e d upwards, strongly
attached with laterai plates, two times longer than broad; posterior processes srnail,
strongly divergent; imer margin of bridge curved upwards, with excavated part in the
middle.
P hall ic complex (except epiphallus) : apodemes short, wide, strongl y divergent, widened
apices; inner projections of cingulum strongly reduced; sheath of penis covenng al1 arch
and apical valves of cingulum, and 60% of apical valves of penis, extending from zygoma
to posterior margin of distal part of rami; rami of cingulum long, wide, wedge shaped,
normally developed with acute distal part; Aedeagus: arch of cingulum long; apical valves
37
of cingulum short, reaching 70% of length of apical valves of penis; apical valves of
penis, with widened anterior margin.
Bohemanella f. fngida (Figure 1 8)
Epiphallus: Size medium; ancorae wide, with obtuse apex, slightly convergent; anterior
processes as long as ancorae, acute apex, slender, convergent; bridge two times longer
than broad; lophi ellipsoid, three times longer than broad; posterior processes wedge
shaped, with acute apex; inner margin of bridge horizontal
Phallic complex (except epiphallus): apodemes slender, elongate, with obtuse converging
ti ps ; imer projections of cingulurn obtuse, strongl y convergent; sheath of p a i s covering
al1 arch of cingulum and apical valves of p a i s and cingulum, extending h m zygoma to
anterior margin of distal end of rami; rami of cingulurn wide in lateral view and well
developed. Aedeag-us: arch of cingulurn wide, obtuse, with extending lower part of
posterior margin; apical valves of cingulurn reduced, with obtuse tip, almost reaching the
distal end of apical valves of penis; apical valves of penis curved upwards apicaily, with
robust rounded part at the anterior part of the ventral margin.
Booneacris g. glaciafis (Figure 19)
Epiphallus: Size small to medium; ancorae long, slender, finger-like, with obtuse tips,
strongly convergent; anterior processes very wide, shorter than ancorae; bridge two times
longer than broad, slightjy excavated .in the center; lophi oval shape, very wide,
articulated with lateral parts, almost perpendicular to the longitudinal axis of the bridge,
two times longer than broad; posterior processes short, rather wide, with acute tip; inner
margin of bridge slightly curved downwards, with small excavated part in the middle.
Phallic complex (except epiphallus): apodemes long and slender, inner projections of
cingulurn slightly reduced, acute, strongly convergent; sheath of penis covering al1 arch
and apical valves of cingulum, and 80% of apical valves of p i s , extending from zygoma
to anterior margin of distal end of Rrn; rami of cingulum elongate, slender, normally
developed. Aedeaw: arch of cingulum extending forward, with denticle in the middle of
the posterior margin; apical valves of cùigulurn long, reaching 60% of the length of apical
valves of penis; apical valves of penis long, elongate, acute apex, distal part sigrnoid
shaped, curved downwards.
Dendroteîîix quercus (Figure 20)
Epiphallus: Size medium; anccirae long, slender, with obtuse tip, strongly convergent;
anterior processes as long as ancorae, with obtuse tip; bridge three times longer than
broad; lophi cylindrical, articulated with lateral plates, t!!ee times longer than broad;
posterior processes short, slender, slightly divergent; imer margin of bridge horizontal,
with excavated part in the rniddle.
Phallic complex (except epiphallus): apodemes elongate, parallel; imer projections of
cingulurn strongly developed, strongly convergent, almost reaching each other's apices;
sheath of penis covering al1 arch and apical valves of cingulum, and 90% of apical valves
of penis, extending from zygoma to apex of distal end of rami; rami of cingulum slender
39
normally developed. Aedeap: arch of cingulum with small denticle extending from the
middle of posterior margin; apical valves of cingulurn acute apex, well developed, almost
reaching the apex of apical valves of penis; apical valves of penis with widened anterior
margin and hump at the middle of ventral margin.
Podisma pedestns (Figure 1 )
Epiphallus: Size medium to large; anwrae smail, with obtuse tip, convergent; anterior
processes as long as ancorae, widened apically; bridge three times longer than wide; lophi
lobiform, four times longer than wide; posterior processes as wide as ançorae , short;
inner margin of bridge curved upwards, with no excavated or projecting parts.
Phallic complex (except epiphallus): apodema slender, long, divergent; imer projections
of cingulurn acute, well developed; sheath of penis covering 90% of apical valves of
cingulum, 80% of apical valves of penis, extending from zygoma to anterior margin of
distal part of Rm; rami of cingulurn well developed, wide in lateral view, expanded
caudally. Aedeagus: arch of cingulum with long obtuse denticle extending from the top of
the posterior margin; apical valves of cingulurn strongly developed, long, almost reaching
the apex of apical valves of penis; apical valves of penis ventral margin curved upwards.
Tribe Prirnnoini
Primnoa primnoa (Figure 2 1 )
Epiphallus: Size large; ancorae wide, short, strongly convergent, with acute tips; anterior
processes very wide, shoner than ancorae, widened tips; bridge very broad, longer than
40
wide; lophi large, articulated with lateral plates, with concave ventral margin; posterior
processes well developed, divergent, obtuse tip; imer margin of bridge curved
downwards.
Phallic complex (except epiphallus): apodemes very wide, long, siightly divergent,
widened tips; inner projections of cingulurn absent; sheath of penis absent; rami of
cingulum short, wide, parallel. Aedeagus: arch of cingulum well developed, apex towards
basal parts of penis; apical valves of cingulum reaching 50% of apical valves of penis;
apical valves of penis very unique, very long, extending laterally from rami, curved
do wnw ards.
Prumnacris raimierensis (Figure 22)
Epiphallus: Size medium; anwrae very smail, wide, with acute tips; anterior processes as
long as ancorae, obtuse; bridge at least two tirnes longer than broad; lophi elongate,
cylinder shaped; posterior processes well developed, long, slender, acute tips, with
strongly convergent apices; imer margin of bridge horizontal, with small excavated part
in the middle.
PhaIIic complex (except epiphallus): apdemes slender, rather short; imer projections of
cingulum small. acute; sheath of penis covering a small ventral part of apical valves of
cingulum, and approximately 60% of the length of apical valves of penis, extending from
middle of rami to the ventral margin of the distal part of rami; rami of cingulum normally
developed, very long. Aedeagus: arch of cingulurn short; apical valves of cingulum as in
figure, reaching apical valves of penis; apical valves of penis wide.
5.1 Range and distribution of phiiiic charaeten in the subfamily Melanopihie
From the illustrations of al1 the species studied, it is evident that the range of
phallic characters in the melanoplines is remarkably wide, as was expected. First, 1 will
refer to the epiphallus, the structure that is described most frequently in previous
taxonomie papers on melanoplines. The epiphallus in the genera studied is bridge shaped,
with the exception of Aptenopedes, where a mernbranous-like interspace separates the
bridge into two parts. In genersl, the epiphalli of the species studied in this work are in
agreement with the previous illustrations (Roberts 1941, Dirsh 1956, Rehn and Randell
1963). Only minute variations from previous work were observeci (usually the shape of
ancorae and anterior processes), basically due to different drawing techniques and angles
used to illustrate the phallic complex.
The lophi are usually cylindrical or upcurved, and rnight project tiom the lateral
plates of the epiphallus or merge with thern. The ancorae are mostly acute, as are the
anterior projections. The imer margin of the bridge can be straight, but it can also have a
small hump or projection in the middle. The posterior projections are often well
developed and divergent.
Concerning the rest of the phallic complex, various characters are found in the
exarnined species. The cingulum is described for the first time in detail for al1 these
genera (when 1 refer to the "cingulum in this section, 1 am not including the arch of
cingulum and the apical valves of cingulum, since they are both discussed later together
with the aedeagus). This structure that bbholds" the aedeagus has not been used in
taxonomy of the melanoplines for severai possible rasons: i) it is very hard to separate it
fiom the aedeagus and study it separately (parts such as the i ~ e r projections break very
easily during dissection), ii)the aedeagus itself provided enough informative characters, so
there was no need to go through M e r dissection, iii) confusion between the dorsd view
with the fiont view might mislead the results. On the other hand, the cingulum could be of
a more significant taxonomie character, especially the absence or presence of imer
projections, as weli as the length and shape of the rami.
As for the aedeagus and its parts, the range of morphology varies greatly, as was
expected. The apical valves of the cingulum could be long, or reduced, or even absent in
some genera. The apical valves of the penis are oflen very elongated and extend further
fiom the sheath. In most of the genera studied, the arch of cingulum is well developed and
quite robust, differing in shape and size. Al1 these differences obsewed in the present
study were expected, especially for the apical parts of the aedagus, where great variation
occurs even between species of the same genus.
On the other hand, it is essential to mention that the basal parts of the aedeagus
have not been fkquently used in the taxonomy either of the Acrididae, nor of the whole
order Orthoptera
involves concave
pattern, ii) their
(sensu stricto). This is reasonable due to i) their unique structure
and convex areas that are tremendously difficult to draw in a constant
interspecific variability, and iii) the difficulty in extracting usehl
43
characters fiom them. These drawbacks have been observai in the present study as well,
and therefore the basal parts of the aedeagus are still considered of limited usehlness in
menanopline taxonomy.
Despite this variability of aedegal parts, a constricted point of articulation between
the apical and basal parts of the aedeagus was present in al1 the genera studied. The
present work confinns for the first time the absence of the sigmoid flexure in the North
Arnerican Melanoplinae examined (a character that separateci them fiom Cantatopinae),
since Amedegnato (1 974) only studied îhe South and Central American genera.
In order to visualize the range of rnorphology in the male genitalia of the
subfamily Melanoplinae, Table 4 was constnicted, as a better alternative compared to
iengthy paragraphic descriptions. The tabulation of different structures between taxa was
used by Vickery ( 1 993) and subsequently proposed by Cohn (1 993).
5.2 Phylogenetic interpretations and suggestions for reclassification in the subfady.
In this part of the discussion, I will provide suggestions about the classification of
the melanoplines that can be inferred fiom the genitalic similarities of the examined
genera. These are recommendations based solely on genitalic characters and may not be
sufficient justifications for erection of a new tribe or subtribe. However, the hypotheses
that will be presented may be of value in fbture classifications in the subfamily. The main
purpose of this work is to compare the different genera and to obtain information about
their relationships, and not to reconstruct the phylogeny of the North American
44
melanoplines, a laborious task that still seems "untouchable". This is due to the limited
number of orthopterists cunently involved in melanopline systematics (Dr. Otte in
Philadelphia, and Dr. Cigliano in Argentina), but also acridoid systematics in general
(Otte 1990). Another reason may be the fact that funds for research have been diverted to
other areas of orthopterology.
It was not possible to identifi the general characteristics of each of the tribes of
the Melanoplinae, leaving the tribal classification still in question. As has already been
stated by other systematists, the group is very variable (not only in genitalic characters)
and it is highly probable that several genera may need to be rearranged or that new tribes
and subtribes might be erected. Roberts (194 1) called this subfamily the dumping ground,
Rehn and Randell (1963) indicated that additional tribes may be found essential, Cohn
and Cantrall ( 1974) admitteci that characteristics that differentiate tribes are variable and
overlapping, while Otte (1995) did not even attempt to assign numerous genera to any
tribe due to their equivocal status. The same problem of variation and questionable s t a tu
of many genera was a factor in the present study as well. Phallic structures sometimes
confirmed the previous classifications (the usual case), neveriheless some new questions
arose for several genera of arnbiguous position.
It shoufd also be realized that only a complete cladistic approach could resolve
questions about the phylogenetic relationships between the studied genera. Similarity in
several important structures is probably evidence of a close relationship, but this could
only be confinned afier the characters are polarized and the pleisiomorphic and
apomorphic States are established.
The most remarkable outcome of the cornparison of different genera, is the status
of Aptenopedes (Table 4 W). The questionable position of this genus was mentioned by
Dirsh (1956) who placed it into the ungrouped and uncertain genera of Cantatopinae.
Later, Rehn and Randell (1963), considered it an atypical melanopline, stating that it
differed from other mernbers of the subfamily because of an apparent separation of the
ancorae fiom the bridge of the epiphallus, a feature absent fiom other melanoplines. There
are more unique character States present in this genus, such as the hook-like lophi, and the
membranous-like separation of the bridge. These characters are not f o n d in any other
melanoplines exarnined, they are not species specific and variable (as for example the
apical valves of cingulum and aedeagus), thus they justify the separation of Aptenopedes
from the rest of the melanopline genera. Vickery @monal oommunication) believed that
there are several extemal characters which confinn the atypical s t a t u of Aptenopedes and
its questionable position in the subfamily. The status of the genus could only be resolved
with a cladistic analysis, in order to test whether this genus actually belongs to a different
subfamily, or it simply represents an autapomorphic lineage within the Melanoplinae.
Usefiil information was also obtained for two genera that have a limited
distribution in the eastem United States and Canada, Appalachia and Dendrotettix (Table
4 I, J). Both of them were placed together by Rehn and Randell (1963) in the group
Podismae. Most of the stmctures of the epiphaIlus (ancorae, inner rnargin of bridge,
postenor processes) and of the apical parts of the aedeagus (apical valves and arch of
cingulum, apical valves of penis) show strong similarity, indicating a possible close
relationship of these bvo genera. This was previously highlighted by both morphological
(Rehn & Rehn 1939), and cytological (Fontana and Vickery 1976, Vickery 1989) data.
Podisma and Booneucris (Table 4 H , K) were also c o n s i d d members of the
group Podismae by Rehn and Randell (1963). This was altered by Fontana and Vickery
( 1 976), due to cytological differences of Booneacris from the r a t of the group, and thus it
was placed in a separate subtribe Mirarnellina, together with severai Palearctic podismine
genera. The present study revealed several difierences observed mainly in the epiphallus
(lophi , ancorae, posterior processes), that are however inadequate to con firm any subtribal
position of Booneucris.
The subtribal status of Bohemanella figida (Table 4 M) also rearnins
questionable. No particular character States could support the placement of the genus in
either subtribe of Podismini. The question might be answered after cytological or
rnolecular studies, as Fontana and Vickery ( 1 976) suggested. However, the genus belongs
to the tnbe Podismini, as previously proposed by Vickery (1984) and not to Melanoplini
(as Melanoplus fiigidus) as suggested by Vickery and Kevan ( 1 983) and Otte ( 1 995).
A group of genera (ffvpochlora, Phoetaliotes, Aeoplides. Table 4 U , T, V) share
the absence of apical valves of cingulum, a stnicture present in other Melanoplini.
Additionally, the apical vaives of the penis are very long and curved upwards, the lophi
are cylindrical and protmde fiom the lateral plates, while the posterior processes are
wedge-shaped and strongly divergent. Therefore they could probably be considered as a
different group from the typical Melanoplini, maybe constituting a separate subtribe. A
common ancestor for these genera of western American origin had already been
hypothesized by Rehn (1954). Again, cladistic evidence is needed to support the
monophyly of sister groups.
Conalcaea and Bavttetix (Table 4 E, F) are corredy placed in tribe Conalcaeini.
Their phall ic stnictures show great divergence fiom Melanoplus, whi le several extemal
characters that seemed "unusual" for a member of Melanoplini, forceù Rehn and Randell
(1963) to place them into the "atypical" group. My results (most of the genitalic
characters of these two genera are strongly divergent fiom other melanoplines studied)
agree with those of Cohn and Cantrall (1974), hence the two genera should still be
considered members of Conalcaeini.
Rehn and Rehn (1944) stated that Pmrnnacris and Primnoa (Table 4 C, D) were
very closely related, and they were later placed in the podismine group Primnoae by Rehn
and Randell (1963). Phallic structures studied in this work indiçate a resernblance of
Prurnnacris with Podisma (lophi cylindrical, well developed imer projections of the
cingulum, long rami), rather than with Melanoplus or Primnoa. Thus, it is suggested that
Pnrmnacris should Se placed in Podismini, and not in Primnoini (as in Rehn and Randell
1963) or Melanoplini (Otte 1995). On the other hand, Primnoa seems highly divergent
from the podismine group, with a remarkable (and unusual for a member of the subfamil y
Melanoplinae) prolongation of the apical valves of the penis extending laterally fiom
rami. The shape of the arch of cingulum is also unique, as is the formation of the lophi.
These characters could possibly justify the placement of Primnoa in the separate aibe
Primnoini, as currently accepted.
The tri be Brad ynotini incl udes Bradvnotes, Asemopfus. A rgiacris and
Buckellacris, (Table 4 N, O, A, B), al1 of western North American distribution. It was
impossible to find common phallic characters to support the placement of these genera in
the same tribe, al1 being rather diverse, with no significant similarities. Despite that,
Argiacris seems be closer to Bradynotes than any of the other genera, because of several
genitalic sirnilarities (lophi of the same shape, divergent posterior processes, reduced
apical valves of cingulum, short rami normaily developed). m i s is in agreement with
Hebard ( 1 9 1 8) who noted the resemblance in extemal appearance of these two genera.
In Buckeffacris, the phailic complex is very peculiar for a member of
Melanoplinae. Its apicai valves of penis are very complex, the rami are unique in their
shape as viewed dorsdly and the cingulum as a whole does not resemble any other genus
studied. 1 wouid suggest that Buckellacris should be removed fiom Bradynotini and
placed in a separate tribe. However, only cladistic analysis could resolve the questionable
position of this genus more objectively, by detennining the polarity of the characters in
the subfamily.
The classification is also questionable for several genera that belong to the tribes
Dactylotini and Dichroplini, namely Dactyiotum, Hespercltettix and Dichropius (Table 4
G, P, K). When Rehn and Randell (1963) gave the general characteristics of Dactylotini
(including Dichropius), the genus Dactyiotum differed in numerous extemal characters
fiom the rest of the tribe. Additionally, the only genitalic character that separated
Dactylotini fiom
the similarity of
Melanoplini was the reduced posterior processes of the epiphallus, while
Dichroplus with Mefanoplus (Table 4 Q, R) was also highlighted. The
49
results fkom rny study, reveal no particular character that could reliably differentiate
Dactylotini fiom Melanoplini. In my opinion, genera Dactylotum and Hespero~tix
should probably constitute a subhibe (Dactylotina) of Melanoplini, while Dichropfus and
some allied South American genera (such as Ronderosia, Atrachelacrïs, Scotussa and
Leiotettix, illustrated in Cigliano 1997) should compose another subtribe (Dichroplina) of
Melanoplini.
5.3 Future research
It should be obvious fiom the above discussion that Mer research in various
aspects of melanopline systematics is grealy needed. There is no doubt about the
necessity of a revision in the subfamily, based both on extemal and genitalic features. The
application of modem cladistic approaches that would analyze and establish the
phylogenetic relationships in this group is the only secure and objective way to proceed
fiom now on, leading to a more objective classification. Otte (personal communication)
has already started to revise the genus Melanoplus, while attempts were made by
Ronderos and Cigliano (1991) and Cigliano (1997) to analyze the phylogeny of some
South Arnerican melanopline genera using cladistics. Hopefully, the present study will
help to stimulate more intensive study of this group of grasshoppers, this time with the
incorporation of techniques of phylogenetic systematics (cladistics).
Despite numerous controversies (e.g. Kluge 1983, Sibley and Ahlquist 1987,
Wagele & Wetzel 1994, Lamboy 1994), most systematists agree that that integration of
50
both molecular and morphological data will maxirnize the validity and usefulness of
phylogenetic hypotheses (Hillis, 1987, Hoy 1994, Wagele & Wetzel 1994, Moritz and
Hillis 1996). Chapco (personal communication) has already started to investigate
relationships between melanopline genera, after atîempting to reclassiQ the whole
orthopteroid superorder (Kupenis and Chapco 1996).
To m l v e al1 phylogenetic problems of such a diverse and neglected group as the
melanoplines, the appropriaîe start wodd be to "abandon names, as classically applied,
and regroup the genera based on their shared, derived characters. This will be an
enornus task, but it is necessary" (Nickle and Naskrecki, 1997). Nickle and Naskrecki
were r e f e g only to katydids and crickets but their staternent is applicable for the
phylogeny of many orthopteroid groups including the melanoplines. The puzzle is huge,
but we should begin putting the pieces together.
6. CONCLUSIONS
From the study of the phallic complex of 22 selected genera of Melanoplinae, the
following results could be inferred:
i) there is a great variation in the intemal male genitalia between the genera of the
subfarnily
ii) the epiphatlus and the apical parts of the aedeagus are the most important taxonomie
characters in this group
iii) the cingulum, a previously neglected (as a whole) structure, might sometimes provide
usefiil information, especially its rami and imer projections
iv) Aptenopedes is highly divergent fiom the rest of the Melanoplinae
v) Hypochlora, Phoefaliofes and Aeoplides may constitute of a subtribe Melanoplini
vi) Subtribal classification of Podisrna, Dendrotetrix. Appalachia and Booneucris is still
uncertain, with a strong resernblance of the three first genera
vii) BohemaneZla resembles the podismine genera, but its subtribal classification is also
questionable.
viii) Conalcaea and Borytetix should remain in Conalcaeini
ix) There were no phallic characters strong enough to support the differentiation of
Dactyiotum and Hesperotettix (both in Dactylotini) from Melanoplini
x) Dichroplus and its allied South American genera should probably consist of a subtribe
of Melanoplini, rather than a single separate tribe
xi) Prumnacris shares more similar phallic characters with the podismines as compared
with Prirnnoa, thus its removal h m Primnoini could be suggested.
xii) no signifiant similarities were found in phallic characters to support the placement of
several western American genera (Bradynotes, BuckeIIacris, Argiocris, Asemoplus) in
a single tribe Bradynotini. In fact, Buckellacris seerns to be highiy divergent fiom the
other studied melanopline genera.
xiii) Cldistic analyses that wouM test the conpence between morphological and
molecular data are considerd asential for the reconstruction of the phylogenetic
relationships between the melanopline genera.
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FIGURES AND TABLES
Figure 1. Phaltic parts of Podisma pedestris. A. Dorsal view of
epiphallus, B. Dorsal view of phallic cornplex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum. E. Lateral view of
aedeagus
Abbreviations: A: anwrae, Ant: anterior processes, Apd: apodemes of
cingulum, Arc: arch of cingulum, Avc: apical valves of cingulum,
Avp, apical valves of penis, B: bridge, Imb: imer margin of
bridge, Bp: basal parts of penis, Ejd: ejaculatory duct, Ejs;
ejaculatory sac, Gpr: gonopore process, L: lophi, Pp: posterior
processes, Rm: rami of cingulum, Sh: sheath of penis, Sps:
spermatophore sac
lmb
Arc
Avc
Avp
Ejs E
Figure 2. Phallic parts of Agriacris rehni. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphailus), D. Frontal view of cingulurn, E. Lateral view of
aedeagus
Figure 3. Phallic parts of Asemoplus monfanus. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphaltus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulwn, E. Lateral view of
aedeagus
Figure 4. Phallic parts of Bradjmotes o. obesa. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Laterai view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. L a t d view of
aedeagus
Figure S. Phallic parts of Buckeiiucris c. chilcotinae. A. Dorsal view
of epiphallus, B. Dorsal view of phailic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 6. Phallic parts of Butyrertix h hymphreysii A. Dorsal view
of epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Front view of cingulum, E. Lateral view of
aedeagus
Figure 7. Phallic parts of Conalcuea miguelimta. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 8. Phallic parts of Dacfylotum bicoiorpictum. A. Dorsal view
of epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Laterd view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 9. Phallic parts of Hesperotetrlx viridis pratensis. A. Dorsal
view of epiphallus, B. Dorsal view of phallic compiex (without
epiphallus), C. Lateral view of phallic cornplex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 10. Phallic parts of Dichroplus elongatus. A. Dorsai view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), Ce Lateral view of phallic complex (without
epiphallus), De Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 11. Phallic parts of Aeoplides t. turbulli. A. Dorsal view of
epiphallus, B. Dorsal Mew of phallic complex (without
epiphailus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulwn, E. Lateral view of
aedeagus
Figure 12. Phallic parts of Apienopedes sphenarioides. A. Dorsal
view of epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagu
Figure 13. Phallic parts of H'hlora dba. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 14. Phallic parts of Mefanopfus femurnrbrum. A. Dorsal view
of epiphallus, B. Dorsal view of phallic complex (without
zpiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulurn, E. Lateral view of
aedeagus
Figure 15. Phallic parts of Paroxya cfmliger A. Donal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phaliic cornplex (without
epiphallus), D. Frontal view of cingulurn, E. Lateral view of
aedeagus
Figure 16. Phallic parts of Phoetaliotes nebrascemis. A. Dorsal view
of epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Laterai view of
aedeagus
Figure 17. Phallic parts of Appalachia arcana. A. Dorsal view of
epiphailus, B. Dorsal view of phailic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphailus), D. Frontal view of cingulurn, E. Lateral view of
aedeagus
Figure 18. Phallic parts of Bohemanella f: fiigida A. Dorsal view of
epiphallus, B. Dorsal view of phallic cornplex (without
epiphallus), C. Lateral view of phallic cornplex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 19. Phallic parts of Booneacris g. glacialis A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 20. Phallic parts of Dendrotettix quercus. A. Dorsal view of
epiphalIus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic cornplex (without
epiphallus), D. Frontal view of cingulum, E. Lateral view of
aedeagus
Figure 21. Phallic parts of Primnoa primnoa. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phalhc complex (without
epiphallus), D. Frontal view of cingulum, E, Lateral view of
aedeagus
Figure 22. Phallic parts of Prumnacris rainierensis. A. Dorsal view of
epiphallus, B. Dorsal view of phallic complex (without
epiphallus), C. Lateral view of phallic complex (without
epiphallus), D. Frontal view of cingulurn, E. Lateral view of
aedeagus
Table 1. Classification of the subfarnily Melanoplinae, as proposed
by Rehn and Randell (1963). Genera in upper case lettering are
those studied in the present work. Note that today, the "groups"
are considerd tribes (Vickay 1997)
i io
Melanoplini Dacty lotiai Podismini
PAROXYA
Chioroplus
Eotettix
MELANOPL us HYPOCHLORQ
PHOETA LIOTES
Phaulotettix
Cephdotettir
Philocleon
Sinaloa
Phaedrorettix
Pedies
Ne caxacr is
Amical
Aidemona
Agroecotettix
Oedaleonotus
AEOP LIDES
BAR YlTETUI
CONALCAEA
APTENOPEDES
HESP EROTEclTJX
P oecilotettix
Campylacantha
Oedemem
P aratyZotropidia
Paraidernona
DfCHROPL US
Chibh4cris
DACTYLOTUM
Dasyscirtus
Perikerus
Aztecacris
Amical
Meridacris
Gymnoscirteres
Grour, Podisma
Micropodismu
Zybovskya
BOONRCRIS
DENLCROTEmLX
APPALACW
Eirenephilw
Ognevia
PODISMA
B O H E W E L L A
gr ou^ Primnoae
P R I W O A
Prumnacris
Gro up Miramellue
hdopodisma
Oropodismu
Cop hopodisma
Epipodisma
Miramella
Chortopodisrna
Purapodisma
Sinopodisma
gr ou^ Bradvnotae
Hypsa onia
He bardacris
ARGL4CNS
BUCKELLACRlS
Kingdonella
ASEMOPL US
BRQDYNOTES
Atyical
Odonropodisma
Pseudopodisma
Yunnanucris
Niitahcris
Pseudoprumna
Table 2. List of genera and exemplar species studied in the present
work
GENUS SPECIES* STUDIED TYPE SPECIES
Argiacris Hebard Argiacris rehni Hebard studicd
Asemoplus Scudder
Brad'vnotes Scudder
BuckeIlacris Rehn & Rthn
Baqvettir Scudder
Conalcaea Scudder
Dac~lotum Charpentier
Hesperotetrir Scudder
3ichroplus SW
Aeoplides Caudell
Aptenopedes Scudder
lfvpochlora Brunner
Mefanoplus StAi
Paropa Scudder
Phoetaliores Scudder
Appalachia Rehn & Rehn
Bohemanella Rammc
Booneacris Rehn & Randcll
Dendroretrü Pac kard
Podkma Berthold
Prirnnoa Fischer- Waldhein
Pnrrnnacris Rehn & Rchn
Asemoplus montanus Bnuier
Brdvnotes o. obesa niornas
Buckelfacris c. chilcotinae Hebard
Ba~tetrix h. humphrqvsii Thomas
Conalcaea miguelinara Scudder
Dac~lotum bicolor pictum Thomas
Hesperorettù viridis pratensis Scudder
Dichroplrrs elongatus Giglio-Tos
Aeoplides t. rurbulli îhomas
Aptenopedes spirenurioides Scudder
Hvpochlora alba (Dodge)
Mefanoplus femumbrum (De Geer)
Paroqa clavuliger (Audinet-Serville)
Phoeialiores nebrascemis (niornas)
Appalachia arcana Hubbell & Cantrdl
Bohemanella J fngida Boheman
BooneacrrS g. glacialis Scudder
Dendroretiix quercus Packard
Podkma pedestris Linnaeus
Primnoa primnoa Fischer-Waldhein
Pmmnacris rainierensis Caudell
studid
studied
studied
BatyrterLx crasus Scudder
studitd
sîudied
siudied
Acridium arrogans Stal
Pezotettù chenopodii CaudcIl
studied
sîudied
sîudied
studied
sîudied
Appalachia hebardi Rehn & Rehn
studied
studied
s tudied
studied
siudied
studied
*or subspecies
Table 3. Tenninology used by different authors for the genitalia of
Melanoplinae
Structure I Walkn (1922) Dirsh (1956)
apophyses du pont endapophyses endapophyses apodemes
arcb of cingulum t-- -
arch of dorsal
valves
arch of cingulum bridge connecting
dorsal lobe with phalIoireme sclerites
arc h
apical valves of
cingulum
basal parts of pcnis
dorsal valves of dorsal valves of
aedeagus
endophallic plaies
aedeagus
endophallic plates basal valves of
penis
rami of cingulum
valves superieures endoparameres
rami of cingulum r- valves inferieures rami rami of latcral plates in the rami of cingulum
endapophyses proximal part of the
dorsal lobe
oukr membrane of
distal part of dorsal
sheath of penis I outer membrane of
dorsal and ventral
lobes
--
sheath of aedeagus sheath of pcnis
lobe of aedeagus
apicrl valves of posterior ventral ventral valves of
aedeagus or
aedeagal valves
apical valves of
penis
cc toparameres
arch of endapophyses
apical process of
aedeagus
pont posterieur zygoma zygoma endapophysal arch
of tpipbrllus
bridge I lophi
posterior projections t-
dccurved spines of
pseudostemite
Snodgrass ( 1935)
postconiua or processes
of ihe pseudostemite
anterior processes
transverse bar of
the pseudostemite
dorsal lobes of
preudostemite
bridge
posterior processes of
the epiphallus
Roberts (1 94 1)
ancorae
bridge
lophi
Dirsh ( 1 956)
ancorae
-- -
anterior
projections of
epiphallus
bridgc
lophi
posterior
projections
Table 4. Comparative table of the intemal male genitalia of the
melanopline genera studied. Al1 figures are drawn to the sarne
sale
- -
Genus Epiphdlus Ectophallur Cingulum
Q Dichopfyt
R Mclonophs
S Puraxya
T Phoetaiiotes
U Hypochiora
v A eoplides
W Aptenopedes
