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 National Library Bibliothèque nationale 1+1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Sewices senrices bibliographiques 395 Wellington Street 395. rue Wellington OmwaON KlA ON1 OnawaON K1AW Canada CaMda
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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 mle 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 & Dr 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 of the material for my study. Moreover 1 would like to extend my gratitude to the Academy of Natural Sciences of Philadelphia and iis 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.
Apm 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 fady in Greece, for supponing me both financially and psychologically
during al1 this time 1 was away fiom home. TABLE OF CONTENTS
Page
* - .USTRACT II RESUME
DEDICATION
ACKNOWLEDGMENTS
TABLE OF CONTENTS
LIST OF FIGWS
LIST OF TABLES
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
... Vlll LIST OF FIGURES
Figure Page
1. Intemal male genitalia of Podisma pedesrris 67
2. Intemal male genitdia of A rgiacris rehni 69
3. Inted male genitalia of Asemopfus montanus 71
4 Intemal male genitaiia of Bradjnotes o. obesu 73
5. Internal male genitaiia of Buckeflacrisc. chifcofinae 75
6. Internal male genitaiia of Barytetfix h. hymphreysii 77
7. Internai male genitalin of Conalcaea miguelinata 79
8. Intemal male genitaiia of Dac~lotumbicolor picium 8 1
9. Intemal male genitalia of Hesperofettix virirüs praiensis 83
10. Internal male genitaiia of Dichropftis elongatus 85
11. Internai male genitalia of Aeoplides t. tumbulli 87
12. Int emal male genitalia of Aptenopedes sphenarioides 89
13. Internal male genitaiia of Hypochfora albû 9 1
14. Internai male genitaiia of Melanopius femzrrmbrz~m 93
15. Internal male genitaiia of Paroxya clavuliger 95
16. Internal male genitalia of Phoetaliotes nebrascer~sis 97
17. Internai male genitalia of Appafachia urcana 99
18. Internal male genitaiia of BohemanellaJ fi.isda 101
19. Internal male genitalia of Booneacris g. gfucialis 103
ix Figure Page
20 Intemal male genitalia of Dendroretrix queras 105
2 1 Internal male genitalia of Primnoa primnoa 1 O7
22 Internal male genitalia of Pmmnacris rainierensis 1O9 LIST OF TABLES
Table Page
1. Classification of the subfamily Melanoplinae, as proposed by Rehn and
Randel1 ( 1 963). I Il
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 1974). This is relative1y 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 1974, 1977, Amedegnato and Descarnps 1979, Ronderos and Cigliano 1990,
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 subsequently followed by Otte ( 1995) and Vickery ( 1997) ] 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 of this project are to: i. Describe in detail the morphology of the interna1 male genitalia of selected North
American genera of 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 subfamil 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 increases. Rehn and Randell (1 963) suggested separation of the subfamily into 3 tribes; Melanoplini, Podismini and Dactylotini. (Table 1)- In Rehn and
Randell 's ( 1 963) work, the characteristics that were used to differentiate the tribes were
4 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 imply that they might not reflect the real phylogenetic histoq of the species; indeed 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 hm other centers of radiation (Vickery 1989). It is believed that these two genera are "sister- genera", wry close1y 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 (19 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 distinctivel 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 ( 193 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 ( 19 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
1935: 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 differences of the subfamilies, new additional 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 fmale 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 (1963) 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 ( 194 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 exclusive1y responsible for the reproductive isolation of the species involved. They took 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 in North Amencan Melanoplinae will definitely increase the number of species and 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. 64x4938 (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 Gig 1io-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- 1961 (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)
Melanoplusfernmurrubrum (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- 1966 (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
USA, 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- 19 10, (N. Ikonikov) ( 1 male, ANSP)
PrumnacriS rainierensis Caudell
1) USA, WA, Mount Rainier, Pierce Co., Camp of 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 ( 1941 ) 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 glas 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. 33 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 hm 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 1stabdominal segment. The phallic organ represents the distal end of the ejaculatory duct and could be divided to two main parts: the aedeagus (Figure
1B,C,E and the cingulum (Figure 1B-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 1A), 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 penetrate the female genital chamber. The epiphallus is dso a significant structure in 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; der 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, vqthick 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 cwed 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 pais 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 pais 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 epiphallus) : 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 hmthe 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, slightly convergent apical1y; 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 pais 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~ermargin of bndge horizontal, with minute denticle in the middle.
Phallic complex (except epiphallus): apodemes slender, moderately long; inner projections of cinylum obtuse, wide; sheath of penis small, covering 70% of 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 12)
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, strongly divergent; inner margin of bridge slightl y curved upwards.
Phallic complex (except epiphallus): apodemes very long and slender, converging apically; inner projections of cingulurn absent; sheath of pais 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 pais 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 pais covexing arch of cingulum, 50%1 of 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, cwed upwards.
Tribe Podismini
Appalachia orcana (Figure 17)
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 cwed 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 18)
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, strongly convergent; sheath of pais covering al1 arch of cingulum and apical valves of pais and cingulum, extending hmzygoma 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 pis, 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 downw 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~erprojections break very easily during dissection), ii)the aedeagus itself provided enough informative characters, so there was no need to go through Merdissection, 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 (sensu stricto). This is reasonable due to i) their unique structure involves concave and convex areas that are tremendously difficult to draw in a constant pattern, ii) their 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 ( 1993) 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 statu 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 fond 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 statu 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 considd members of the group Podismae by Rehn and Randell (1963). This was altered by Fontana and Vickery
( 1976), due to cytological differences of Booneacris from the rat 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 confirm 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 (1976) 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 ( 1983) and Otte ( 1995).
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 subfamily
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). mis 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 Melanoplini was the reduced posterior processes of the epiphallus, while the similarity of 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 mlveal1 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 refeg 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, Phoefaliofesand 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 hmPrimnoini 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
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15: 1-76. 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. Latd 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 Mefanopfusfemurnrbrum. 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)
iio Melanoplini Dacty lotiai Podismini
PAROXYA HESP EROTEclTJX Grour, Podisma gr ou^ Bradvnotae Chioroplus Poecilotettix Micropodismu Hypsa onia Eotettix Campylacantha Zybovskya He bardacris MELANOPL us Oedemem BOONRCRIS ARGL4CNS HYPOCHLORQ ParatyZotropidia DENLCROTEmLX BUCKELLACRlS PHOETALIOTES Paraidernona APPALACW Kingdonella Phaulotettix DfCHROPL US Eirenephilw ASEMOPL US Cephdotettir Chibh4cris Ognevia BRQDYNOTES Philocleon DACTYLOTUM PODISMA Sinaloa Dasyscirtus BOHEWELLA Atyical Phaedrorettix Perikerus Odonropodisma Pedies Aztecacris gr ou^ Primnoae Pseudopodisma Ne caxacr is PRIWOA Yunnanucris Prumnacris Niitahcris Amical Amical Pseudoprumna Aidemona Meridacris Agroecotettix Gymnoscirteres Group Miramellue Oedaleonotus hdopodisma AEOP LIDES Oropodismu BAR YlTETUI Cophopodisma CONALCAEA Epipodisma APTENOPEDES Miramella Chortopodisrna Purapodisma Sinopodisma 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 Asemoplus montanus Bnuier studid Brad'vnotes Scudder Brdvnotes o. obesa niornas studied BuckeIlacris Rehn & Rthn Buckelfacrisc. chilcotinae Hebard studied Baqvettir Scudder Ba~tetrixh. humphrqvsii Thomas BatyrterLx crasus Scudder Conalcaea Scudder Conalcaea miguelinara Scudder studitd Dac~lotumCharpentier Dac~lotumbicolor pictum Thomas sîudied Hesperotetrir Scudder Hesperorettù viridis pratensis Scudder siudied 3ichroplus SW Dichroplrrs elongatus Giglio-Tos Acridium arrogans Stal Aeoplides Caudell Aeoplides t. rurbulli îhomas Pezotettù chenopodii CaudcIl Aptenopedes Scudder Aptenopedes spirenurioides Scudder studied lfvpochlora Brunner Hvpochlora alba (Dodge) sîudied Mefanoplus StAi Mefanoplusfemumbrum (De Geer) sîudied Paropa Scudder Paroqa clavuliger (Audinet-Serville) studied Phoetaliores Scudder Phoeialiores nebrascemis (niornas) sîudied Appalachia Rehn & Rehn Appalachia arcana Hubbell & Cantrdl Appalachia hebardi Rehn & Rehn Bohemanella Rammc Bohemanella J fngida Boheman studied Booneacris Rehn & Randcll BooneacrrS g. glacialis Scudder studied Dendroretrü Pac kard Dendroretiix quercus Packard studied Podkma Berthold Podkma pedestris Linnaeus studied Prirnnoa Fischer- Waldhein Primnoa primnoa Fischer-Waldhein siudied Pnrrnnacris Rehn & Rchn Pmmnacris rainierensis Caudell studied
*or subspecies Table 3. Tenninology used by different authors for the genitalia of
Melanoplinae IStructure Walkn (1922) Dirsh (1956) apophyses du pont endapophyses endapophyses apodemes - arcb of cingulum bridge connecting arch of dorsal arch of cingulum dorsal lobe with phalIoireme sclerites valves arc h t-- t-- apical valves of dorsal valves of dorsal valves of cingulum aedeagus aedeagus basal parts of pcnis valves superieures endoparameres endophallic plates endophallic plaies basal valves of penis rami of cingulum valves inferieures rami rami of latcral plates in the rami of cingulum rami of cingulum endapophyses proximal part of the dorsal lobe
-- r-sheath of penis outer membrane of oukr membrane of sheath of aedeagus sheath of pcnis dorsal and ventral distal part of dorsal lobes lobe of aedeagus Iapicrl valves of cc toparameres apical valves of posterior ventral ventral valves of apical process of aedeagus or penis aedeagus aedeagal valves pont posterieur arch of endapophyses endapophysal arch zygoma zygoma Snodgrass ( 1935) Roberts (1 94 1) Dirsh (1 956)
dccurved spines of anterior processes ancorae ancorae pseudostemite
-- - anterior of tpipbrllus projections of epiphallus bridge transverse bar of bridge bridge bridgc I the pseudostemite lophi postconiua or processes dorsal lobes of posterior processes of lophi lophi of ihe pseudostemite preudostemite the epiphallus posterior projections posterior t- 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