A GENERIC LEVEL REVISION AND CLADISTIC ANALYSIS OF THE OF THE WORLD (, LYGAEIDAE, )

B. JANE HARRINGTON

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el A GENERIC LEVEL REVISION AND CLADISTIC ANALYSIS OF THE MYODOCHINI OF THE WORLD (HEMIPTERA, LYGAEIDAE, RHYPAROCHROMINAE)

B. JANE HARRINGTON Assistant Professor, Department ofEntomology University of Wisconsin

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 167: ARTICLE 2 NEW YORK: 1980 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 167, article 2, pages 45-116, figures 1-103 Issued October 28, 1980 Price: $4.10 a copy

ISSN 0003-0090

Copyright ©) American Museum of Natural History 1980 CONTENTS Abstract 49 Introduction. 49 Acknowledgments.50 Systematic History of the Tribe Myodochini .51 Materials and Methods .52 Character Analysis.54 Phylogeny.61 Distribution and Zoogeography .64 Classification.67 Key to the Genera of the Tribe Myodochini .69 Generic Diagnoses and Descriptions.77 Literature Cited.110

ABSTRACT The present study examines the rhyparochro- talisi; Paraparomius lateralis, P. leptopoides; mine tribe Myodochini at the generic level on a limbatus; albomaeulata, P. world basis. A key to 56 genera is provided and caeca, P. confusa, P. dissimilis, P. foeda, P. for- diagnoses and brief summary descriptions of sa- mosa, P. fusicornis, P. intercisa, P. minima, P. lient morphological features are given for all gen- oblonga, P. plebeius, P. tumens; Pseudopachy- era. brachius basalis, P. capicolus, P. nesovinctus, P. Thirteen new genera (Ashlockaria, Bacaceph- pacificus, P. reductus, P. undulatus, P. vinctus; alus, Ereminellus, Froeschneria, Humilocoris, Pseudopamera ater, P. insititius, P. setosa; Pseu- Megacholula, , Paracholula, Para- doparomius linearis; Remaudiereana africana, R. paromius, Pseudoparomius, Slaterobius, Stalar- andrewsi, R. annulipes, R. boninensis, R. casta- ia, and Stridulocoris) and the new species Me- nea, R. flavipes, R. horvathi, R. inornatus, R. gacholula englemani are described. kydippe, R. nigra, R. nigriceps, R. noctuabundus, The following new synonymies are made: Ex- R. octonotata, R. puberulus, R. robustus, R. sid- with Prytanes; Neocattarus with nicus, R. sobrina, R. tibialis; Slaterobius insignis, Cholula; Sphaerobius with Erlacda; and Sphaero- S. quadristriatus; Stalaria ferruginosus, S. kis- bius gracilis with Erlacda arhaphaeoides. seis, S. nysias; Stigmatonotum afrus, S. genicu- New combinations include: Ashlockaria sob- latus; Stridulocoris gracilis. rius; Bacacephalus globiceps; Cholula firmus, C. A cladistic analysis of the myodochine genera irrorandus, C. maculatus, C. parvus, C. vigens; is presented with emphasis on the valuable char- Ereminellus arizonensis; Froeschneria infumatus, acter system of the male genitalia. Plesiomorphic F. multispinus, F. oblitus, F. piligerus, F. vicin- and apomorphic states are designated for all char- alis; Horridipamera bergrothi, H. ebenaui, H. in- acters employed, and only synapomorphic char- conspicuus, H. perlongus, H. pullatus, H. rusti- acter states are employed to infer relationship and cus, H. spinicrus, H. subsericeus; Humilocoris to construct a generic cladogram for the tribe cephalotes; occultus; Neopamera al- Myodochini. bocinctus, N. bilobata, N. brachialis, N. costalis, The phylogeny shown in the cladogram is N. crassicornis, N. honduranus, N. insularis, N. viewed against the current known distributions of intermedius, N. mumfordi, N. neotropicalis, N. the various living myodochine species and used paganus, N. platanus, N. procerulus, N. recinc- to propose a broad zoogeographic hypothesis tus, N. serripes, N. sororculus, N. tineodes, N. about the origins of the tribe and major mono- tuberculatus, N. vicarius, N. vividus; Orthaea phyletic assemblages within the Myodochini. Al- consuta, 0. procinctus; Pamerana punctulatus, most total lack of transoceanic sister groups at P. scotti, P. sinae; Paracholula thoracicus, P. the generic level suggests a postGondwanaland vegetus; Paraeucosmetus albofasciatus, P. cras- origin and evolution for the tribe, and the distri- siceps, P. gemmatus, P. harimaensis, P. mala- butional patterns also attest to the isolated evo- yus, P. mimulus, P. nervosus, P. sladeni, P. vi- lution of two major lineages in North America.

INTRODUCTION The Myodochini, with 56 genera and more Some myodochine genera seek seeds still in than 260 species, is the largest tribe in the the seed head, others in the ground nests of lygaeid subfamily Rhyparochrominae. Mem- rodents, and yet others in bird droppings on bers occur in all six major zoogeographic re- leaves. The majority of the species, how- gions and representatives also are known ever, are ground bugs. There they can usu- from several remote Pacific Oceanic islands. ally be found in the litter layer, feeding on All members of the tribe whose feeding fallen seeds within the natural seed shadow habits are known are seed-predators. In fact, of a plant. Sweet (1964) has studied the bi- except for the blood-feeding tribe Cleradini, ology of certain Nearctic members in detail this specialized phytophagous habit is char- and reports some seed-defense behavior. acteristic for the subfamily (Sweet, 1964). Most of the myodochini are small, dull- 49 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

colored, often cryptic and many are tory, Smithsonian Institution) for allowing not generally collected except by specialists. me to study (and subsequently borrow) ma- Despite the large number of taxa described terial and type specimens from the collec- since Stal's (1874) key to genera, the group tions of these museums and for all their co- had not attracted much taxonomic interest, operation and kindness during my visits. beyond the alpha level and a few generic re- I also thank the following individuals for visions, until quite recently, when Malipatil their contributions to this work: Dr. H. D. (1978) published a revision of the tribe for Engleman for his hospitality and aid in col- the Australian region. lecting in Panama; Dr. Dingemans-Bakels, My initial involvement with the tribe was Naturrhistorisch Museum, Maastricht, the an effort to revise the large cosmopolitan ge- Netherlands, for kindly searching (without nus . Upon discovering that success) the Wasmann Collection for the Pachybrachius was highly polyphyletic, I type of Fontejanus wasmanni; Drs. C. W. saw the necessity for generic level work be- Schaefer and F. A. Streams, for critically fore any meaningful alpha could reading the manuscript; and Dr. Peter D. be done. The present resulting work is the Ashlock, University of Kansas, for the loan first modern worldwide examination of the of Kodachrome transparencies of several tribe in its entirety. Emphasis has been type specimens and for personal discussion placed on delimiting the genera and propos- and guidance in cladistic thinking and the art ing a generic phylogeny. of constructing cladograms. Special thanks are due to Dr. James A. ACKNOWLEDGMENTS Slater, my advisor at the University of Con- necticut, for the generous loan of specimens I appreciate the loan of numerous speci- and literature from his personal collection mens for study by the following individuals and library, for the loan of the dorsal view and institutions: Mr. S. L. Szerlip (Univer- of triguttatus drawn by Mr. Ste- sity of California, Berkeley), Dr. N. M. An- phen Thurston (University of Connecticut) dersen (Universitetets Zoologiske Museum, and for his advice, example and unfailing in- Copenhagen), Dr. P. D. Ashlock (Snow Mu- terest and encouragement. seum, University of Kansas), Dr. T. R. I especially appreciate the efforts of the Yonke (University of Missouri), Dr. J. D. late Mrs. D. B. Wilcox whose painstaking Lattin (Oregon State University), Dr. Johann checking of the key was an important service Becker and Jose C. M. Carvalho (Museau but whose friendship, personal warmth, and Nacional, Rio de Janeiro), Drs. J. C. Schaff- good-humored support through all phases of ner and M. H. Sweet (Texas A & M Uni- this study were of greatest value. versity), Dr. F. Fernandez-Yepez (Univer- An important collecting and field study trip sidad Central de Venezuela, Maracay) and to several islands in the West Indies was gen- Drs. P. D. Ashlock, H. D. Engleman, J. A. erously supported by National Science Slater and M. H. Sweet private collections. number GB 27162, Special thanks are due to Drs. R. T. Schuh Foundation Travel Grant and P. Wygodzinsky (American Museum and NSF additionally funded a collecting trip Natural History), Dr. W. J. Knight and Mr. to Panama. A University of Connecticut Re- W. R. Dolling (British Museum (Natural His- search Council grant allowed me to travel to, tory)), Dr. G. E. Wallace (Carnegie Mu- and partially supported an extended study of seum) and Drs. R. C. Froeschner and J. L. type material at, the British Museum (Nat- Herring (National Museum of Natural His- ural History). 1980 HARRINGTON: MYODOCHINI 51

SYSTEMATIC HISTORY OF THE TRIBE MYODOCHINI Although there were earlier groupings, as chiefly of a complex of Australian taxa which noted in Slater (1964a), the Myodochini were Gross (1962) had designated as "Aberrant first defined and named by Stal (1872) as the Australian Brachypterous Myodochine division Myodocharia. Even at that early Bugs," to a new tribe , and sug- date, Stal clearly recognized the positions of gested the probable origin of the Myodochini the abdominal spiracles as good indicators of from a Neotropical udeocorine stock. Mem- higher group relationships within the bers of the Myodochini are distinguished subfamily Rhyparochrominae. Yet, Breddin from members of the tribe Udeocorini by the (1907) was the first to actually characterize lack of inner laterotergites. the tribe Myodochini (which he referred to Although many myodochine species were as the Pameroidea) as having abdominal spi- described by such early workers as Lin- racles II through IV dorsal. Currently, this naeus, Fabricius, Hahn, Dallas, Herrich- derived pattern of abdominal spiracles re- Schaeffer, Say, Signoret, Guerin, Fieber, mains the most reliable character for distin- Uhler, Baerensprung, Motschulsky, and guishing the tribe. Walker, the first major study of the group as Since Stal's early definition, the major no- a whole was Stal's key to the genera in 1874. menclatural problem for the group has been Working with the large and diverse Neotrop- the status of the name for the type genus, ical fauna, Distant (1880-1893) described nu- . Slater, Barber and Sailer (1959, merous species and nearly one-third of the 1961) have discussed this elaborate case in genera for the tribe. Around the turn of the detail. Their 1961 appeal to the International century, additional descriptions were con- Commission on Zoological Nomenclature tributed by European workers such as Ber- led, in 1963, to Opinion 669 which relegated groth, Reuter, Horvath, Scott, Kirkaldy, and Myodochus Olivier, 1811 to the Official List Breddin. of Rejected and Invalid Generic Names in Early in the twentieth century, Barber be- Zoology; designated Myodocha Latreille, gan a more thorough analysis, primarily of 1810 as the official name of the genus, plac- the Nearctic fauna. He described several ing it on the Official List of Generic Names new genera and species, producing many in Zoology; recognized Myodocha serripes publications on the group, the most notable Olivier, 1811 as the type species of the genus of which are his three careful generic revi- and placed that species name on the Official sions (1921, 1928a, 1953). List of Specific Names in Zoology; and Contemporary work on the Myodochini placed the family-group name Myodochini has been published by Ashlock, China, Hob- Stal, 1872 (correction of Myodocharia Stal, erlandt, Scudder, Slater, Sweet, and Usin- 1872 by Van Duzee, 1916) on the Official List ger; new genera are still being described. Of of Family-Group Names in Zoology. the total of 56 myodochine genera recog- Numerous authors, including Scudder nized in this study, four were described be- (1957), have followed Puton (1879) in calling fore 1851, 14 in the interval of 1851-1900, 10 the group the Plociomerini, and, as noted from 1901-1925, three from 1926-1950 and above, Breddin (1907) used the name Pa- six from 1951-1976. Quite recently, Malipatil meroidea. Yet, both are clearly junior syn- (1978) revised the tribe for the Australian re- onyms of Myodochini, Plociomera, in fact, gion and described nine new genera, five of being a misspelling of Ptochiomera Say (see them for previously described species. How- Barber, 1928). Scudder (1957) reduced the ever, only six of those nine genera are in- taxon to a subtribe of his expanded Rhypar- cluded among the 56 genera dealt with in this ochromini, but Slater and Sweet (1961) re- study (see classification section for expla- turned it to tribal status. nation). In the present study 13 new genera Sweet (1967) removed a group of Austra- are recognized, 12 of them for previously lian and Neotropical genera, composed described species. 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

In view of the amazing diversity of the most certainly remain to be discovered. tribe in the Neotropics, additional taxa al- MATERIALS AND METHODS Determined material of some species has the Myodochini. (The key to genera which been available for all but two of the 40 pre- is given is simply a device for identification viously described myodochine genera listed or a classificatory aid, and makes no attempt in Slater (1964a). I was able to examine over to reflect phylogeny.) 80 percent of the approximately 260 named This work departs from strict cladistics as species. advocated by Hennig in three basic facets: Dried and alcoholic specimens were ex- 1. Sister groups are not afforded the same amined with a binocular dissecting micro- absolute rank, as they would be in a cladistic scope under powers ranging from lOx to system. With the Myodochini, adherence to 80x. Measurements were taken by means of such a scheme of classification would de- a calibrated ocular micrometer and drawings mand that, if the monotypic genus Kolene- were made to scale with the use of an ocular trus is recognized as a genus, the entire rest grid. However, because of great size dispar- of the world's myodochine species must be ity among the genera, the drawings were not lumped together in a taxon also of generic all made at the same magnification. In de- rank. Clearly, this is an extreme case, but it scriptions the Villalobos color chart (Palmer, does serve to hold up to question the basic 1962) was used as a standard. cladistic tenet of equal rank for sister groups. The male genital capsule was removed 2. Hierarchical rank has not been based on from relaxed specimens with jewelers for- actual or inferred geological ages of the taxa ceps; cleared for approximately three min- in this study and even most cladistic studies utes in a hot potassium hydroxide solution have not adhered to this original Hennigian and dissected in distilled water in porcelain stricture. spot plates. The dissected material was 3. Paraphyletic genera are recognized and transferred through another distilled water named. As stressed by Hennig (1966), only bath, a bath of 10 percent acetic acid and a synapomorphous, or shared derived, char- solution of 70 percent ethanol (five minutes acters are admissible as evidence of evolu- each) and placed in glycerin for further study tionary relationship while symplesiomorphy, and drawing. Once fully examined, the gen- or sharing of primitive traits, does not imply italia were stored in glycerin in microvials relationship. pinned with the proper specimens. In cases According to the specifications of Hennig where tribal status was questionable, the ab- (1965), a monophyletic group is based on domen was removed and similarly cleared in synapomorphy, a polyphyletic group on con- potassium hydroxide to reveal spiracular po- vergence, and a paraphyletic group on sym- sitions and the presence or absence of inner plesiomorphy. Yet, even with reference to laterotergites. Hennig's own diagrams (fig. 2), I suggest that After study of the various genera, alter- this is not really the case with regard to par- nate character conditions or states were de- aphyly. Paraphyletic taxa are not deliberate- noted as plesiomorphic (primitive, ancestral, ly recognized and established on the basis of or generalized) or apomorphic (derived, de- symplesiomorphy. They indicate, instead, scendant, or specialized). Often the validity incomplete knowledge and are often an in- of these inferences was enhanced by estab- dication of how thoroughly a group has been lishing congruence with other character dis- studied. In the cladogram (fig. 103) of my- tributions, as recommended by Hennig odochine genera, for example, Cholula is a (1966). Synapomorphic, or shared derived paraphyletic genus. The lack of synapomor- characters were then utilized to construct the phy (not the use of symplesiomorphy) for cladogram of figure 103, which presents a recognizing this genus has been indicated by proposed phylogeny, or possible course of a dotted line. Cholula could be thought of as evolutionary events, at the generic level for being synonymous with ancestor 5. 1980 HARRINGTON: MYODOCHINI 53

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The monophyletic assemblage removed synapomorphy is lacking but there is not from Cholula (ancestor 5) and rendering contrary evidence indicating convergence Cholula (ancestor 5) paraphyletic is the ge- (polyphyly), the practice of naming paraphy- nus Stridulocoris. Stridulocoris is defined on letic genera should be more generally ac- the complex and firm synapomorphic state cepted. Later more detailed study of the of having an abdominal stridulatory appara- group may reveal other character systems tus (figs. 31, 32). This is an important unique- that present the lacking synapomorphy(ies), ly derived character, as the two species of or perhaps these derived character states Stridulocoris are the only myodochines of may be behavioral, physiological, etc., and the large and diverse fauna with male geni- go undetected indefinitely without rigorous talic Type I to possess stridulatory struc- biological studies not typically undertaken in tures. Here then, the cladist's stricture a taxon as vast as the Insecta. In some cases against recognizing paraphyletic genera it is possible that autapomorphic derivations seems to demand that evidence actually be may even have obliterated evidence of the discarded or denied. With stress on the im- lacking synapomorphy. Such possibilities portance of synapomorphy, the cladist's ap- must have been in the mind of Farris (1974) proach has reached the shaky stance of "if when he stated "it is probable that there ex- you don't have two, you must disregard ist monophyletic groups in the conventional one." sense which possess no single distinguishing At the particular juncture represented by character state." hypothetical ancestor 5, I can either recog- It should also be remembered that any nize the paraphyletic genus Cholula, against cladogram (be the lines solid or dotted) is the dictates of cladistics, or suppress the evi- merely a hypothesis. The demand is placed dence of the strong synapomorphy defining not on how a hypothesis is formulated but Stridulocoris, against the dictates of accept- rather on its internal consistency, explana- ed scientific procedure and common sense. tory power, predictive power, and potential I have chosen the former path. On such a to be falsified (Popper, 1959). Subject to basis, other paraphyletic genera, all indicat- these criteria, paraphyletic taxa should be ed by dotted lines, are recognized and named just as admissible as monophyletic ones, and in this study of the Myodochini. On this a solid line should have no more inherent same basis, I would argue that, when a firm validity than a dotted one. CHARACTER ANALYSIS Generally, in the absence of other evi- as ancestral to the Myodochini, have been dence, a character state occurring broadly considered plesiomorphic in the Myodochi- within the tribe Myodochini was assumed to ni. The Udeocorini is the only rhyparo- be plesiomorphic, while the alternative con- chromine tribe, other than the Myodochini, dition, occurring in a limited number of taxa, whose members have abdominal spiracles 2- was viewed as apomorphic. Yet, outgroup 4 dorsal and thus it must be considered the comparison has also been very important in sister group of the Myodochini. Udeocorine determining the plesiomorphic or apomorph- genera surveyed in this study for external ic states of certain characters. A condition character conditions and dissected for geni- such as pronotal carination found widely talic comparison include Astemoplitus, throughout all the other tribes of the subfam- Euander, Fontejus, Neosuris, Porander, ily Rhyparochrominae was considered ple- Serranegra, Telocoris, and Udeocoris. siomorphic, whether common or rare in the As in many other groups, the scler- Myodochini. The alternative condition or otized parts of the male genitalia have pro- character state was designated apomorphic. vided an excellent character system which I In particular, character states found in the have emphasized in this cladistic analysis. Udeocorini, the tribe Sweet (1967) suggested The decision of apomorphic vs. plesio- 1980 HARRINGTON: MYODOCHINI 55 morphic for various characters of the exter- this genus is recognized, but, currently it is nal morphology has often been based on cor- retained in the group with Type I male gen- relation with the patterns established by the italia. genitalic characters, as well as by the criteria The following three phallic types have all indicated above. arisen independently from Type I by my The following characters have been found current interpretation, and these three sep- to be important to the classification of the arate apomorphic states are not to be con- Myodochini. Additional minor characters, strued as a series of character states arising often applicable to only a single genus, are from or grading into each other. discussed separately in the individual generic (Type II). Ten genera have a Type II phal- diagnoses where any suspected instances of lus (fig. 10). In this apomorphic phallic parallelism or convergence are also dis- condition, presented by Bacacephalus, Eu- cussed. cosmetus, Horridipamera, Mimobius, MALE GENITALIA: Terminology and in- Paraeucosmetus, Paraparomius, Paromius, terpretation of the various elements of the , Pseudoparomius and lygaeid phallus were discussed in detail by Togo, the vesica is adorned by sawtoothed Ashlock (1957), who noted the diversity of rows of minute, closely spaced spines. Hold- myodochine phallic types. The myodochine ing sclerites appear to be lacking entirely and phallus presents an excellent character sys- the wings of the well-developed sperm res- tem for analyzing relationships and drawing ervoir are large and broad, rolling under lat- zoogeographic inferences for the tribe. Al- erally in a scroll-like fashion. Two large, though there are a few distinctive autapo- heavily sclerotized, hollow spines (in some morphic variations in detail (described in the cases unequal with one bifid and in Mimo- individual generic diagnoses), the 56 myodo- bius branched and multiheaded) appear on chine genera fall into four basic phallic types: the conjunctiva very close to its vesical junc- (Type I). The plesiomorphic condition ture. In the uninflated phallus these large found throughout the Udeocorini examined spines lie dorsad of the sperm reservoir. and known to occur in other rhyparochro- (Type III). The apomorphic Type III phal- mine tribes, is present in the myodochine lus (fig. 11) is characterized by a conjunctiva genera Aegyptocoris, Caenopamera, Carpi- adorned by many spines, some being lis, Cholula, Ereminellus, Fontathanus, grouped in rows or patches. The sperm res- Henicorthaea, Humilocoris, Kolenetrus, ervoir is reduced and elongate with narrow, Megacholula, Megapamera, Pachybrach- straplike, anteriorly directed wings. Holding ius, Pamerarma, Pamarana, Pamerapa, sclerites are lacking and there are no spines Paracholula, Prytanes, Ptochiomera, Sis- on the tubular vesica. The five genera in this amnes, Remaudiereana, Stalaria, Stigma- group, Froeschneria, Ligyrocoris, Peri- tonotum, Stridulocoris, Suffenus, and Valo- genes, Slaterobius, and Zeridoneus are pri- netus. This condition (fig. 9) is characterized marily a Nearctic element, although most of by a well-developed sperm reservoir with the species of Froeschneria occur in South anterolaterally directed wings; long, slender, America. parenthesis-like holding sclerites; and usu- (Type IV). Sixteen myodochine genera ally unadorned conjunctiva and vesica. Fon- have a Type IV phallus (fig. 12). This phallic tathanus, Henicorthaea, Megapamera, type is not so morphologically distinctive as Type II and Type III. The sperm reservoir Pachybrachius, Pamerana, Pamerapa, and is well developed with broad wings. Holding Stalaria show some autapomorphic varia- sclerites are typically present, though short- tions of the conjunctiva but retain the char- ened and characteristically flattened and acteristic parenthesis-like holding sclerites of broadened distally. The vesica is unadorned the Type I phallus. Pamerarma, with una- and the tiny spines which occur (but rarely) dorned conjunctiva and vesica, totally lacks on the conjunctiva are very unlike and prob- holding sclerites. The genitalic uniqueness of ably not homologous with those of Type III. 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

Afrovertanus, Ashlockaria, Catenes, Cne- rior pronotal collar can be traced in the modus, Distingphyses, Erlacda, Heraeus, Myodochini. Kolenetrus has the plesio- Myodocha, Neopamera, Orthaea, Pephy- morphic condition (fig. 58), with no indica- sena, Pseudocnemodus, Pseudopamera, tion of a collar dorsally on the anterior pro- Tenuicoris, Xenydrium, and Zeropamera notal lobe. share phallic Type IV. In Cholula (fig. 50), Megacholula (fig. 62) The dorsal aperture of the myodochine py- and Suffenus (fig. 55) a narrow collar can be gophore or male genital capsule has four discerned only medially. character conditions: In Paracholula (fig. 64) and Stridulocoris (A). The posterior edge of this opening is (fig. 59) a very narrow collar is present very sharp (fig. 4) in Aegyptocoris, Afrover- across the entire anterior portion of the pro- tanus, Cholula, Henicorthaea, Kolenetrus, notum. Megacholula, Paracholula, and Stridulocor- Aegyptocoris (fig. 43) and Fontathanus is. This sharp-edged condition, common in (fig. 46) show only a slight indication of a the Udeocorini, is considered the plesio- collar. morphic character state for the myodo- Caenopamera (fig. 51), Carpilis (fig. 57), chines. Ereminellus (fig. 65), Henicorthaea (fig. 54), (B). Almost half of the myodochine genera Humilocoris (fig. 60), Prytanes (fig. 49), Pto- have an apomorphic condition that can be chiomera (fig. 56), Stigmatonotum (fig. 63), best described as "subsharp" (fig. 5). In- and Valonetus (fig. 61) have a distinct collar cluded in this group are Caenopamera, Car- region noticeable in the lateral pronotal mar- pilis, Ereminellus, Fontathanus, Pachy- gin from above but not posteriorly demarked brachius, Prytanes, Pseudocnemodus, by a groove from the rest of the anterior pro- Ptochiomera, , Stigmatonotum, notal lobe. and Xenydrium, and all genera with phallic All other myodochine genera have a very Types II and III. distinct collar which is demarked posteriorly (C). With the above-mentioned exceptions by a fine, linelike groove. In some cases, at ofAfrovertanus, Pseudocnemodus, and Xen- high magnification, this line appears as a ydrium, genera with phallic Type IV have the fused row of shining punctures sunk between posterior margin of the aperture to the male the adjoining pruinose areas of the collar and genital capsule broadly rounded (fig. 6). This the rest of the anterior pronotal lobe. condition is considered apomorphic relative Pamerarma (fig. 99), Pamerapa (fig. 44), to the subsharp state. Remaudiereana (fig. 52), and Stalaria (fig. (D). The remaining apomorphic condition 47) are related on the basis of a distinctive, is exhibited by only a few Old World genera. narrow, ringlike collar which is totally im- In Pamerapa, Pamerarma, Remaudiereana, punctate and very deeply demarked poste- and Stalaria the posterior margin of the cap- riorly. sule aperture is fairly broadly rounded but Other myodochine collar variations in- with a noticeable median depression (fig. 7). clude lateral spines on the collar of Xeny- This depression is very deep and distinctive drium (fig. 92), the blunt "beginnings" of in the autapomorphic state ofPamerana (fig. similar but independently derived spines 8). seen on the collar of some specimens of Togo The claspers or parameres of the Myodo- (fig. 76), the peculiar median V-shape to the chini show minor variation from species to collar of Paromius (fig. 66), and the dorsally species. They have limited value above the narrow but ventrally very broad and ante- specific level and have not been used in this riorly produced collar of Heraeus (fig. 87). generic analysis. CLAVAL PUNCTATION: Three regular PRONOTAL COLLAR: A variety of char- rows of punctures (fig. 19) represent the ple- acter states involving the origin, develop- siomorphic condition of claval punctation. ment and posterior demarcation of an ante- This character state is found in Aegyptocor- 1980 HARRINGTON: MYODOCHINI 57 is, Erlacda, Kolenetrus, Paracholula, Pry- Tenuicoris (all ant mimics) the median carina tanes, Ptochiomera, Sisamnes, Slaterobius, is considerably prolonged beyond the buc- Suffenus, and Valonetus. Although the three cular juncture. rows of claval punctures presented by Sla- Caenopamera, Carpilis, Ereminellus, Pry- terobius and Erlacda (both with derived tanes, Ptochiomera, Sisamnes, and Valone- phallic types) may represent a retention of tus with the synapomorphic condition of a the plesiomorphic condition, they might also grooved midventral surface of the head con- be viewed as evolutionary reversions possi- taining a U-shaped buccular juncture close bly associated with the brachyptery of these to the labial insertion (fig. 80) are essentially two genera. a Nearctic element. The tiny Old World ant The state considered apomorphic, on the mimic Aegyptocoris is unique with a basis of correlation with the phallic types, U-shaped juncture close around the base of typically occurs as four or more rows of cla- the labium but with the ventral surface of the val punctures with only the two rows nearest head not grooved. the corium very precisely aligned (fig. 17). EYES AND OCELLI: Most myodochine This is the condition found in most myodo- genera have essentially round eyes. This is chine genera and probably has arisen more the plesiomorphic condition. than once. Some genera such as Caenopa- Elongate, oval eyes are found in Disting- mera, Carpilis, Ereminellus, and Ligyrocoris physes (fig. 89), Pephysena (fig. 86), Tenui- are transitional, with but a few scattered coris (fig. 96), and in some species of Eucos- punctures added between two of the three metus (fig. 71), and Paraeucosmetus (fig. rows of the plesiomorphic type (fig. 18). 100). This condition has arisen more than BUCCULAE: Three character states in- once and is probably related to the alteration volving the bucculae and ventral surface of of head shape involved in the ant mimicry of the head are found in the tribe Myodochini. species of these genera. Cholula, Kolenetrus, Megacholula, Para- The small, monotypic genus Valonetus has cholula, Stridulocoris, and Suffenus have the peculiar substalked, beadlike eyes and ocelli bucculae prolonged over most of the ventral (fig. 61). surface of the head and enclosing a gular alone among the myodochines area. Their ultimate juncture posteriorly is lacks ocelli (fig. 98). In other myodochine rounded or U-shaped (fig. 79). This, the ple- genera the position of the ocelli relative to siomorphic state, is found throughout the the eyes has been used as a key character. Udeocorini as well as in these "primitive" But this is a morphological area that has been myodochines with the plesiomorphic phallic altered repeatedly in the evolution of the Type I. Included within this group is Para- tribe and thus has not been used to infer re- cholula with a peculiar development of the lationship of genera. bucculae (fig. 78). STRIDULATORY MECHANISMS: Sweet Most myodochine genera exhibit an apo- (1964) has discussed the stridulatory behav- morphic condition little differentiated from ior of a number of North American Myodo- the plesiomorphic state described above but chini as part of a courtship ritual. Ashlock still clearly recognizable (fig. 81). In this and Lattin (1963) described in detail the mi- apomorphic state, the buccular juncture is crotextures of lygaeid stridulitra and the lo- not rounded but V-shaped and is often pro- cation and conformation of plectra, including longed posteriorly as a slight midventral ca- those of the three types of stridulatory mech- rina. The bucculae typically join mesally at anisms found in the Myodochini. Most my- about the level of the antenniferous tuber- odochines are nonstridulatory. This is the cles, yet, in Erlacda, Henicorthaea, Humil- plesiomorphic state. ocoris, Stigmatonotum, and Xenydrium the The monotypic genus Pseudocnemodus is V-shaped juncture occurs closer to the labial characterized by its peculiar autapomorphic insertion. In Distingphyses, Pephysena, and stridulatory apparatus. Arclike, cross-striat- 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 ed areas on the propleuron and the lateral cal in other rhyparochromine tribes, the surface of the head form the stridulitrum (fig. pronotum is dorsoventrally compressed and 39) and the plectrum consists of a row of a slight lateral carina is present on both the tubercles on the proximal third of the fore anterior and posterior lobes, though often it femur (fig. 40). is only clearly apparent in the area of the Other myodochine stridulatory mecha- transverse impression between the lobes (fig. nisms have lunate, filelike stridulitra laterally 22). Among the myodochines, this condition on the abdomen. Afrovertanus and Erlacda is present in Ereminellus in the southwestern have in common a short stridulitrum restrict- United States, Megacholula in the Neotrop- ed to sterna II and III (first two visible) (figs. ics, and Suffenus in the Old World. 33, 28). The plectrum for this type of strid- The Nearctic monotypic genus Kolenetrus ulatory apparatus consists of two or three has a very similar compressed condition that large, chisel-like projections (figs. 34, 29) at may be termed "subcarinate." Among the the base of the hind femur. A mechanism Neotropical relatives of Megacholula, Chol- similar to that of Afrovertanus and Erlacda ula has a compressed pronotum but the ca- involves a larger stridulitrum which extends rination is visible only on the posterior lobe; additionally onto sternum IV (fig. 35), where- Paracholula and Stridulocoris are less com- as numerous small tubercles arranged in an pressed and show no real carination. Pto- irregular row or a scattered field on the basal chiomera and Sisamnes, which are related to portion of the hind femur form the plectrum Ereminellus on the basis of a midventral (fig. 36). Froeschneria, Ligyrocoris, and head groove, have a distinct carina on the Pseudopamera clearly present the latter type posterior pronotal lobe but a rounded ante- of stridulatory apparatus. It is also present rior pronotal lobe. but less apparent in Ashlockaria and Slater- The remaining myodochine genera lack obius. any indication of the plesiomorphic carina The related genera and Zerido- and have the pronotum apomorphically neus, which clearly belong in the Ligyrocoris rounded laterally on both lobes. complex on the basis of male genitalia, have ACETABULUM: The acetabulum for the no such apparatus readily visible, but Sweet middle leg is formed by elements of both the (1964) noted what might be construed as meso- and metathorax. In the plesiomorphic stridulatory behavior in the courtship ritual state (present in most rhyparochromine of Zeridoneus. Courtship behavior has not tribes, including the Udeocorini and a ma- been observed for Perigenes. While the lack jority of the Myodochini) a narrow portion of stridulatory structures in Perigenes and of the metepisternum curves forward and Zeridoneus might be explained as a second- touches the mesepisternum so that the mes- ary loss of such a mechanism within this lin- epimeron is enclosed between them (fig. 37). eage, a more parsimonious explanation sug- In the alternative (apomorphic) condition, gested by the stridulatory behavior is that the the mesepimeron is prolonged and "emer- morphological adaptations for stridulation gent" or not enclosed by the two episternal just never developed in these two genera. parts (fig. 38). This derived state appears to A rather diffuse abdominal stridulitrum have arisen independently at least four times (fig. 31), sometimes not readily apparent, has within the tribe Myodochini. In all instances evolved independently in the monotypic ge- it is found in long-legged very rapid running nus Stridulocoris. The plectrum in this case forms, being present in Afrovertanus, Me- is a scattered field of very minute tubercles gapamera, Pamerana, Slaterobius, and the on the base of the hind femur (fig. 32). large pruinose Neotropical element with PRONOTUM: A progression of character Type IV male genitalia that includes Ca- states can be traced in the myodochine tenes, Heraeus, Myodocha, Neopamera, pronotum (often best observed in lateral Orthaea, Pephysena, and Tenuicoris. view). In the plesiomorphic condition, typi- WING POLYMORPHISM: In the tribe My- 1980 HARRINGTON: MYODOCHINI 59 odochini wing polymorphism or wing reduc- surface texture) area of the metapleuron and tion has obviously evolved repeatedly and is a narrow posterior band on the mesopleuron not employed in relating genera in this study. form an evaporative area for the metathora- At least 15 genera, with members from each cic scent gland of the adult myodochine. In of the four phallic types, are known to ex- the plesiomorphic condition this evaporative hibit brachyptery or submacroptery. Often area is broad, covering a large portion of the such wing reduction is accompanied by a glo- metapleuron (fig. 42). bose anterior pronotal lobe. A few myodochine genera, apparently in CORIAL PUNCTATION: Several of the my- conjunction with a derived shining body sur- odochine genera considered "primitive" on face, have a narrow or reduced evaporative the basis of Type I plesiomorphic male gen- area (fig. 41). This appears as a repeated italia show scattered punctures along the trend or series of convergences in myodo- membranal margin of the corium. This hem- chine evolution. It occurs in such disparate elytral condition is also found in various genera as Aegyptocoris, Afrovertanus, Ko- udeocorines. lenetrus, Pseudopamera, and the Nearctic The myodochines show two opposing de- phallic Type I group of Prytanes and its rel- rived states of this character. In most my- atives. odochine genera all evidence of this specific ANT MIMICRY: Ant mimicry is common row of corial punctation is lost and the mem- among the Myodochini, presumably as a pro- branal margin is very slightly elevated and tection from vertebrate predators. Sweet smoothly rounded (fig. 20). In Stigmatono- (1964) discussed the nature of ant mimicry in tum of the Old World and a Nearctic group, the Rhyparochrominae in some detail. including Carpilis, Prytanes, Ptochiomera, In several genera, of which Xenydrium and Sisamnes, the opposite is true. These (fig. 92) is a striking example, this mimicry genera have a distinctive row of membranal is readily apparent in museum specimens. margin punctures on the corium (fig. 21). But observation of other living myodochines This extensive development of corial punc- in the field has revealed that such genera as tation is also viewed as apomorphic. Slaterobius (fig. 69), Cnemodus (fig. 98), and STERNAL SCALLOPING: In several rhy- many others which may not appear particu- parochromine tribes, including the Udeocor- larly antlike are actually excellent mimics by ini, the anterior margin of abdominal ster- virtue of their behavior. The interrupted, zig- num II (first visible) is scalloped. In some zag pattern of their rapid running makes instances, this pattern is also repeated in the them almost indistinguishable from the ants lateral portion of the juncture of sterna III common in the same habitats. and IV. Most myodochine genera appear to Ant mimicry, obviously a matter of re- have lost such scalloping. However, certain peated convergence in the tribe Myodochini, genera clearly retain it. Traces of such a pat- cannot be used alone to infer relationship. I tern (fig. 27) can be seen at the anterior edge have also tried to avoid the use of any mor- of the sternum just under the metapleuron in phological features that might be construed Fontathanus, Henicorthaea, Kolenetrus, to be a consequence of ant mimicry. In- Stigmatonotum, Suffenus and the Nearctic stances of ant mimicry are noted in the in- complex of genera related to Prytanes. dividual generic diagnoses. In the apomorphic state exhibited by Chol- HEAD ELONGATION: The evolutionary ula, Paracholula and Stridulocoris, the scal- significance or environmental pressures that loping is accentuated and quite clearly ap- lead to an elongation of the head in the My- parent, being prolonged as a longitudinal odochini is a long-standing puzzle. Blatchley cellular division of much of segment II (fig. (1926) compared Myodocha serripes (fig. 97) 26). to the peculiar carabid beetle Casnonia SCENT GLAND EVAPORATIVE AREA: The pennsylvanica (Linnaeus) and suggested that rugose shagreened (see discussion of body the slender necks of both the bug and the 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

beetle had developed by their reaching into fore leg is heavily spined. Rows of large crevices after prey. In such a situation, the spines can be detected along the inner and mandibles of the beetle would be efficient, outer edges and numerous small spines fill but the manipulation of a long labium seems the area between these rows, especially to- virtually impossible and Sweet (1960, 1964) ward the distal end of the femur (fig. 13). has conclusively demonstrated the seed- In the apomorphic condition shared by feeding habit of the myodochini. Pamerapa, Pamerarma, and Remaudi- Myodocha (fig. 97) presents the extreme ereana (fig. 14), there is no outer row of in head elongation for the tribe, but Aegyp- spines; the inner row is present; and the tocoris (fig. 43), Afrovertanus (fig. 88), Dis- small spines, seen between the rows in the tingphyses (fig. 89), Erlacda (fig. 85), Her- plesiomorphic state, are lacking, except for aeus (fig. 87), Pephysena (fig. 86), Tenuicoris one which is greatly enlarged and typically (fig. 96), and Xenydrium (fig. 92) also show falls about mid-length on the femur. a distinct neck region to the head. Addition- Afrovertanus, Catenes, Ereminellus, Hu- ally, Ashlockaria (fig. 90), Catenes (fig. 93), milocoris, Kolenetrus, Ligyrocoris, Peri- Cnemodus (fig. 98), Horridipamera (fig. 74), genes, Prytanes, Ptochiomera, Slaterobius, Megapamera (fig. 82), Orthaea (fig. 91), Pa- Stalaria, Stigmatonotum, Suffenus, Humil- merana (fig. 53), Pamerarma (fig. 99), Par- ocoris, and Zeridoneus species have spines aparomius (fig. 73), Pseudocnemodus (fig. only along the inner edge of the fore femur 84), Pseudopamera (fig. 94), Togo (fig. 76), (fig. 16). Zeropamera (fig. 95), and some species of The remaining myodochine genera have Eucosmetus (fig. 71) and Paraeucosmetus double-ranked spines (fig. 15). The minor (fig. 100) have elongate heads that vary in middle spines of what I consider the plesio- length behind the eyes but lack a distinct morphic condition have been lost and only neck region. the rows on the inner and outer edges of the Fifteen of the genera showing the apomor- fore femur are retained. phy of head elongation are of phallic Type These various patterns of fore femoral IV. But Aegyptocoris, Megapamera, Pa- spine reduction have occurred independent- merana, and Pamerarma of Type I and Eu- ly. The type, number, and arrangement of cosmetus, Horridipamera, Paraeucosmetus, spines often are characteristic for individual Paraparomius, and Togo with Type II male genera. genitalia indicate that head elongation is a In addition to spined femora, males of phenomenon that has been arrived at inde- some myodochine genera possess tibial pendently several times in the evolution of spines. This appears to be yet another ex- the Myodochini. This trend is also repeated ample of repeated independent derivation. A outside of the Myodochini in such rhyparo- male fore tibial spine is found in the Nearctic chromine tribes as the Cleradini and the group with a Type I phallus in Carpilis, . Caenopamera, and Prytanes. The Old World FORE LEG ARMATURE: The armature of genus Megapamera also has a Type I phallus the myodochine fore leg is a complex but and a spined fore tibia, but, in this instance, very useful character system for cladistic there are multiple spines along the length of analysis. The details of autapomorphic con- the tibia in both sexes, which are clearly dis- ditions of single genera are discussed in the tinctive and derived independent of the sin- individual generic diagnoses. gle spine condition in the other three genera There are four basic conditions of the fore of phallic Type I. femur in the tribe. In the plesiomorphic Eucosmetus, Horridipamera, Paraparom- state, evidenced by several udeocorine gen- ius, and Togo, all with a Type II phallus, era and Aegyptocoris, Cholula, Erlacda, have a spined male fore tibia. None of the Paracholula, and Stridulocoris among the genera with Type III male genitalia have tib- Myodochini, the entire ventral surface of the ial spines, but a spined fore tibia is found in 1980 HARRINGTON: MYODOCHINI 61 the Type IV genera Ashlockaria, Cnemodus, in some Udeocorini and some indication of Erlacda, Pseudocnemodus, Xenydrium, Zer- this texture is seen in several myodochine opamera, and in some species of Pseudo- genera with plesiomorphic Type I male gen- pamera. italia, it is an extensive shagreening of the TEXTURE OF BODY SURFACE: The ple- entire body surface, such as that presented siomorphic condition of the myodochine by Ashlockaria, Caenopamera, Cnemodus, body surface, like that of the Udeocorini, is Fontathanus, Megapamera, Pseudocnemo- strongly and densely punctate. Punctation is dus, and Zeropamera that I consider the apparent dorsally on the head and both pro- apomorphic state. notal lobes. The lateral and ventral thoracic With regard to pruinosity, a shining or surfaces are also punctured, including the non-pruinose body surface, seen in only cer- pro-, meso- and metacetabula. An impunc- tain myodochine genera, is considered the tate head, an impunctate anterior pronotal apomorphic condition. This derivation oc- lobe and the impunctate collars of Pamera- curs in several separate lineages within the pa, Pamerarma, and Remaudiereana are tribe. Certain distinctive patterns of pruinos- successive derivations or apomorphic con- ity characterizing single genera are discussed ditions reached in the course of myodochine in the individual generic diagnoses. evolution. The extremely accentuated punc- ANTENNAE: Accompanying a general tation of the dorsal body surface seen in trend of increased body size in the Myodo- Cholula and its relatives is viewed as another chini is an increase in the length of the legs apomorphic state. and antennae that has not been used in this In addition to the varying degrees and ex- cladistic analysis. Whether long or short, the tent of punctation discussed above, the my- antennae are typically slender, all segments odochine body surface may be either shining being essentially uniform in diameter. How- or dull in various areas. A dull appearance ever, Carpilis, Ptochiomera, and Sisamnes may be created by a minutely crazed or exhibit markedly thickened antennae that are roughened sharkskinned texture that is re- viewed as apomorphic conditions. In Carpi- ferred to as shagreened or by the presence lis all the antennal segments are enlarged and of a bloom or pruinosity on the surface, barrel-like (fig. 23), whereas Ptochiomera which, at least in the Blissinae, the scanning and Sisamnes share an apomorphic condi- electron microscope has revealed to be cre- tion in which the terminal two segments are ated by minute cell hairs not distinguishable greatly swollen (figs. 24, 25). Field observa- under ordinary magnifications (Slater and tions of these latter two genera have revealed Harrington, 1970). that these heavy antennae are actually used Although small shagreened patches can be for righting an insect which has been turned found on the dorsal surface of the pronotum on its back.

PHYLOGENY The cladogram of figure 103 presents a shared derived characters. Empty inter- phylogeny or most probable course of evo- nodes, where I have been unable to find a lutionary events for the Myodochini based shared derived character, are indicated by on the current level of knowledge arrived at dotted lines and admittedly represent weak in this study. Each number at a "node," or points or at least lack of knowledge for that branching point, represents a hypothetical portion of the evolutionary scheme as do the ancestor. The hatch marks on the "inter- trifurcations from ancestors 2, 10, 26 and 36 nodes," or lines connecting hypothetical and the quadrifurcation from ancestor 13. ancestors, represent synapomorphous or The four major male genitalic types are in- 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 dicated along the bottom of the cladogram. 11-Sisamnes: dorsum with scalelike hairs giving The grid of distributional information given a granular appearance. at the top of the cladogram is explained in 10-Prytanes: EMPTY INTERNODE. the Distribution and Zoogeography section. 2-13: juncture of bucculae V-shaped (fig. 81). 13-Fontathanus: elongate and deeply bilobed A list of the synapomorphic character pronotum (fig. 46). states used to construct the myodochine ge- 13-14: collar broad and apparent, contrastingly neric cladogram of figure 103 follows: pale, but not demarked posteriorly. 1-Kolenetrus: body surface highly shining; evap- 14-Henicorthaea: head very small and declivent orative area reduced (fig. 41). (fig. 54); fore femoral spines exaggerated, con- 1-2: collar indicated medially. trastingly pale with dark apices. 2-3: punctation extreme, deep pits giving a pebbly 14-15: fore femoral spines single ranked. texture; head very broad and flat with eyes ap- 15-Humilocoris: elongate body hairs. pearing to rest on "square-shouldered" prono- 15-Stigmatonotum: characteristic row of corial tum. punctures along membranal margin (fig. 21). 3-Megacholula: head impunctate; sternal scallop- 13-16: collar demarked posteriorly by a linelike ing lost. groove. 3-4: sternal scalloping accentuated and prolonged 16-17: anterior pronotal lobe, save collar, im- as cellular division of sternum II (fig. 26). punctate. 4-Paracholula: scutellar apex elevated, thumb- 17-18: head elongate; mesepimeron emergent (fig. like, pale; bucculae prolonged and exaggerated 38). (fig. 78). 18-Megapamera: extremely large; several fore 4-5: more than three regular rows of claval punc- tibial spines in both sexes. tures (fig. 17). 18-Pamerana: median groove in posterior lip of 5-Cholula: EMPTY INTERNODE. pygophore (fig. 8). 5-Stridulocoris: diffuse abdominal stridulitrum 17-19: general reduction in punctation; collar (fig. 31). ringlike, largely impunctate (fig. 45). 2-6: collar apparent laterally. 19-Pachybrachius: eyes small and head small, 6-Suffenus: pale and extremely minute; fore fe- somewhat elongate behind eyes (fig. 45); minor mur strongly incrassate and with characteristic autapomorphic variation on phallic Type I (see row of fine, closely ranked spines along inner diagnosis). edge. 19-20: ringlike collar entirely impunctate, ex- 6-7: juncture of bucculae U-shaped, near labial tremely narrow; fore femoral spines single insertion (fig. 80). ranked. 7-Aegyptocoris: head prolonged posteriorly in a 20-Stalaria: male genitalia with spatulate multi- stalklike neck (fig. 43); scutellar apex upturned; spined holding sclerites. highly shining dorsally. 20-21: male fore tibia spined. 7-8: ventral surface of head with a broad median 21-22: robust, subovoid body form; enlarged fore groove (fig. 80). femur. 8-Ereminellus: fore femoral spines reduced. 22-Pamerapa: minute spines on conjunctiva. 8-9: lateral margins of anterior pronotal lobe 22-Remaudiereana: curving male fore tibia with rounded. exaggerated spine. 9-Caenopamera: antennae, especially segment II, elongate. 21-Pamerarma: loss of holding sclerites. 9-10: lacking dorsal pronotal pruinosity, often 16-23: phallic Type II (fig. 10): two large, heavily highly shining; reduced evaporative area (fig. sclerotized spines at extreme apex of conjunc- 41). tiva; sawtoothed spiral of minute spines on ves- 10-Valonetus: juga protruding like tiny tusks; ica. substalked, beadlike eyes and ocelli (fig. 61). 23-Mimobius: very swollen ant mimetic head; 10-11: antennae thickened. apical conjunctival spines branched, multi- 11-12: hemelytra pale, unpatterned. headed. 12-Ptochiomera: antennae capitate, segments III 23-24: dorsal lip of pygophore rounded. and IV markedly swollen (fig. 24). 24-Paromius: anterior pronotal lobe markedly 12-Carpilis: antennae clavate, all segments uni- lower than posterior lobe; posterior margin of formly thickened (fig. 23). pronotal collar V-shaped medially (fig. 66). 1980 HARRINGTON: MYODOCHINI 63

24-25: elongation or elaboration of one or both 38-39: stridulatory behavior. apical conjunctival spines. 39-40: abdominal stridulitrum (figs. 33, 28, 35, 25-26: broad head, broader than transverse 31). impression. 40-Pseudopamera: plectrum a field of spines on 26-27: further elongation of both apical conjunc- hind femur (fig. 36). tival spines. 40-41: head elongate, forming a neck; plectrum 27-Pseudoparomius: left apical conjunctival two or three chisel-like spines on hind femur spine bifid. (figs. 29, 34). 27-Pseudopachybrachius: reduced body size; 41-Erlacda: juga forming a marked shelflike pro- largely pale hemelytra. jection above antennal segment I. 26-28: an extra pair of fleshy or sclerotized lobes 41-Afrovertanus: mesepimeron emergent (fig. proximal to the paired apical conjunctival 38). spines. 39-Pseudocnemodus: head and propleural stri- 28-Paraeucosmetus: EMPTY INTERNODE. dulitrum (fig. 39). 28-Bacacephalus: beadlike eyes; swollen ant mi- 38-Xenydrium: spine on lateral margin of collar; metic head; extra conjunctival lobes slender body ant mimetic with abdomen narrowed an- and markedly pigmented or sclerotized. teriorly; lateral margin of head expanded as a 26-29: spined male fore tibia. pseudomandible between eye and antennal in- 29-Eucosmetus: black and white patterned hem- sertion (fig. 92). elytra; markedly unequal apical conjunctival 37-42: body shagreened. spines, the left very elongate, the right short 42-43: multiple male fore tibial spines; long body and broad. hairs. 29-Togo: wing reduction; right apical conjuncti- 43-Ashlockaria: abdominal stridulitrum (fig. 35). val spine bifid. 43-Zeropamera: body and hemelytra unmarked, 25-30: anterior pronotal lobe globose, impunc- uniformly dark chestnut brown. tate, shagreened; transverse impression deeply 42-Cnemodus: lacking ocelli (fig. 98). incised. 36-44: broad head with a flat vertex; preocular 30-Horridipamera: apical conjunctival spines un- portion of head grooved laterally beneath jugal equal, right with a very broad base. ridge. 30-Paraparomius: EMPTY INTERNODE. 44-45: head with a neck; eyes oval. 13-31: departure from Type I male genitalia by 45-46: head heavily punctate with a pebbly tex- reduction of parenthesis-like holding sclerites. ture. 31-32: phallic Type III (fig. 11): elongation and 46-Distingphyses: anterior pronotal lobe shining. reduction of sperm reservoir; conjunctival 46-Pephysena: neck elongate, quite pronounced spines. (fig. 86). 32-33: abdominal stridulitrum (fig. 35). 45-Tenuicoris: mesepimeron emergent (fig. 38); 33-34: fore femoral spines single ranked. head impunctate; anterior pronotal lobe gla- 34-Slaterobius: shining, globose anterior pronotal brous. lobe. 44-Neopamera: EMPTY INTERNODE. 34-Ligyrocoris: EMPTY INTERNODE. 36-47: vertex of head rounded; incipient head 33-Froeschneria: transverse impression deeply elongation. incised (fig. 67). 47-Orthaea: characteristic orange spots along lat- 32-35: characteristic small cap or crown of as- eral margin of posterior pronotal lobe, medially sociated conjunctival spines; sperm reservoir on posterior pronotal lobe and at corial apex; very reduced and elongate. fore femoral spine reduction. 35-Perigenes: clasper reduced, truncate; narrow 47-48: head distinctly elongate. ringlike collar (fig. 72). 48-Catenes: characteristic parallel-sided head 35-Zeridoneus: dark, stout bristles on all tibiae. shape (fig. 93); darkly sclerotized spots on pale 31-36: phallic Type IV (fig. 12); large bulblike femora. sperm reservoir; holding sclerites short and 48-49: neck stalklike. broad or lacking. 49-Heraeus: ventral portion of collar produced 36-37: dorsum shining or subshining, no pruinos- anteriorly. ity; male fore tibia spined. 49-Myodocha: very long neck (fig. 97); head gla- 37-38: dorsum shining. brous. 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 DISTRIBUTION AND ZOOGEOGRAPHY

As stressed by Ball (1975), only once a "generalized tracks," such as those sought phylogeny has been proposed can any at- to establish vacariance patterns in the man- tempt be made at historical biogeography. ner of Croizat, Nelson, and Rosen (1974), are The cladogram of figure 103 is a graphic hy- apparent. In fact, among the 56 genera in this pothesis, proposing a phylogeny or history generic level analysis, there are but three of evolutionary events for the tribe Myodo- genera (Paromius, Pseudopachybrachius, chini. Also included at the top of the clado- and Ligyrocoris) with species in both hemi- gram is a charting of the distributions of con- spheres and only two generic pairs (from temporary myodochine genera. The numbers ancestors 28 and 41) involving an exclusively in the grid at the top of figure 103 indicate transoceanic sister genus. how many species of a given genus are The plesiomorphic genitalic Type I in- known from a particular zoogeographic re- cludes nearly half of the myodochine genera. gion. Twelve Type I genera are recognized in the For the sake of simplicity, Madagascar has New World, whereas 13 different genera been lumped with the Ethiopian region, Ja- with this phallic type are known from the Old pan is included in the Palearctic, and Pacific World. Thus there is not a single Type I ge- islands not clearly associated with either the nus common to both hemispheres. The phy- Australian or Oriental regions are grouped as logeny proposed here does not present any "Pacific Oceanic islands." The character Type I genera with a transoceanic sister-ge- system of the male genitalia has proven most nus. Such negative evidence strongly sug- useful in trying to draw zoogeographic infer- gests a postGondwanaland radiation for the ences about the Myodochini. As previously tribe. discussed, there are four distinct phallic In the Western Hemisphere there are three types identifiable in the tribe (see discussion phyletic elements within the plesiomorphic under character analysis for descriptions and genitalic Type I. Element 1, the Prytanes detail). These four phallic types are indicated group, is a complex of small, short-bodied along the bottom of figure 103. The historical species including Valonetus, Sisamnes, Pto- zoogeographic analysis below follows from chiomera, Ereminellus, Carpilis, Caenopa- the phylogeny depicted in the cladogram mera, and Prytanes. This group is held to- considered in conjunction with current gether by such apomorphic features as a known distributions. reduced evaporative area (fig. 41), a grooved Species of Myodochini are found on all ventral surface of the head and a U-shaped major land masses from Canada to Chile and buccular juncture (fig. 80). It appears to be Argentina, and from Sweden and Siberia to primarily a Nearctic complex, with only South Africa, Madagascar, and Australia. species of Prytanes known to occur in the Members are also known from many remote Neotropics through much of South America. oceanic islands. In fact, initial inspection of Element 2 contains only the monotypic the currently known myodochine distribu- genus Kolenetrus. It is morphologically quite tions shows the perplexing situation of mem- isolated and has a broad Nearctic distribu- bers of all four phallic types occurring in tion which extends down the mountains at both the eastern and western hemispheres. least to Guatemala. Yet a closer examination reveals that the The monophyletic Element 3 includes Myodochini are not really so ubiquitous. In Cholula, Paracholula, Megacholula and fact, the myodochine faunas of the Old Stridulocoris. Although this group is at pres- World and New World are basically distinct ent Neotropical, there is an Oligocene fossil from each other and apparently have evolved from Wyoming assigned to Cholula by Scud- separately for a long period of time, as no der (1890). I have examined this fossil. It is 1980 HARRINGTON: MYODOCHINI 65 poorly preserved but the general appearance are known from many remote oceanic is- is unlike any modern Cholula and I question lands and have probably reached the New this placement. World relatively recently by overwater Among the Old World Type I genera both transport. Suffenus and Aegyptocoris are isolated and Horridipamera and Paraeucosmetus also not closely related to any other living my- occur extensively throughout the Old World odochines. Suffenus is extremely "primi- tropics but no species of either genus are tive," exhibiting plesiomorphic conditions known from the Western Hemisphere. One for almost all characters examined. The ge- species of Paraparomius occurs widely in nus is monotypic and widely distributed Europe; the three species of Togo are en- through the Old World tropics. Aegyptocor- demic to Japan; and Eucosmetus is limited is, with two African species, is a minute ant to the Oriental region. Mimobius was origi- mimic which has many autapomorphic char- nally described from Madagascar. Neither acters associated with mimicry. the type nor any determined material could The remaining 11 Eastern Hemisphere be located. But a Ghanaian series of a tiny genera of Type I share a derived V-shaped ant mimic, assumed to be a Mimobius from buccular juncture (fig. 81). Stigmatonotum comparison with the original description and and Remaudiereana are widely distributed figures, was examined. and have typical Type I genitalia. The other The presence of two Type II genera, Ba- nine genera are more restricted in their dis- cacephalus and Pseudoparomius, in South tributions. Fontathanus and Stalaria are America is puzzling. Yet, in light of the dis- found in tropical Africa; Pachybrachius is tributions presented by the rest of the tribe, essentially a Palearctic isolate; Humilocoris especially the genera with plesiomorphic, is found in the mountains of India; Henicor- Type I, male genitalia, it seems more likely thaea and Pamerapa are confined to the that the Neotropical occurrence of these two Australian region; Pamerana is Oriental; Pa- genera is due to postGondwanaland, over- merarma occurs on several Oriental islands, water introduction of their ancestors than to in the northern Australian region and on is- vicariance. However, only a species-level lands throughout the Pacific; and Megapa- analysis, especially of all the Pacific Ocean mera is known only from the Solomon Is- island members of phallic Type II, can hope lands and New Guinea. Fontathanus, to answer this question. Henicorthaea, and Humilocoris have typical Genitalic Type III seems to be clearly a Type I genitalia, but the other restricted gen- Nearctic component, which arose not later era show minor unique variations in the male than the Miocene (fossil myodochines rec- genitalia (see individual generic diagnoses ognizable as Type III forms are known from for detail). the Miocene fossil shales of Colorado (Scud- Of the 56 genera in the tribe only Parom- der, 1890). The concentration of southwest- ius, and Pseudopachybrachius, both of phal- ern United States species today suggests that lic Type II, have species occurring naturally perhaps this group arose on the Mexican pla- in both the Old World and the New World. teau coincident with the rise of the Madro- These two genera are widespread in both Tertiary Flora, and subsequently spread hemispheres but with a preponderance of northeastward and, to a limited extent (Li- species in the Eastern Hemisphere. A defin- gyrocoris, Froeschneria, and a single ques- itive understanding of the distribution of the tionable species of Perigenes), into the Neo- Type II group requires further species-level tropics. The occurrence of the single, cladistic analysis, yet it is evident that the otherwise Nearctic species Ligyrocoris syl- group's evolution has been primarily in the vestris in Europe is probably due to a recent Old World tropics. Species of the two genera introduction (possibly post-Pleistocene). named above are a dominant grass-feeding In Phallic Type IV the entire assemblage element with strong dispersal ability. They of genera arising from hypothetical ancestor 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

36 form the dominant myodochine group in Cretaceous or at least when Africa and South the Neotropics today. Only a few species of America were still in close enough proximity the major genera Neopamera, Heraeus, and to account for the broad distribution of the Myodocha occur in the eastern Nearctic. plesiomorphic (Type I) genitalic condition in The remaining Type IV myodochines oc- both the Old World and the New World. The cur largely in Central America and the south- relative lack of old stocks in the Palearctic western United States. Yet, one of their and in southern Australia substantiates the number, the monotypic genus Afrovertanus notion that the tribe evolved after the break- presents the greatest irregularity in the dis- up of Pangaea and the beginning of the frag- tributional patterns of the Myodochini. The mentation of Gondwanaland. only specimens examined were from eastern Genitalic Type II evolved subsequently in Africa and there are no other stridulatory the Old World tropics and appears to have myodochines known from the Old World. entered the Western Hemisphere by over- Afrovertanus exhibits certain autapomorphic water sweepstakes colonization. conditions (see generic diagnosis) that seem Type III, as discussed above, is a Nearctic to indicate a relatively long period of evolu- element. Type IV has also evolved in the tionary isolation. It has a superficial habitus New World after considerable drift of South very similar to that of Myodocha but its sis- America from Africa had occurred but when ter group among the living myodochines is those two plates were still close enough to the New World genus Erlacda (South Amer- allow Afrovertanus to reach Africa. While ica and West Indies). the occurrence of Afrovertanus in Africa is Acknowledging that there are many un- highly anomalous in the myodochine distri- described species and possibly even genera butional pattern, similar distributions are in the Myodochini; that the distributions of known in several flowering plant taxa. The some species are very poorly known; and Bromeliaceae, Sapotaceae, and Magacaceae that fossil material is scant, with none known each have great numbers of Neotropical gen- before the Oligocene, the following analysis era and a single genus in Africa (Hutchinson, represents the most probable zoogeographic 1973). Such distributions, in keeping with interpretation for the tribe based on current Darlington's (1957) hypothesis, might be the myodochine distributions and cladistic affin- result of a Gondwanaland development with ities and the increasing firmness of drift the- subsequent extinction in the Old World by ory. more dominant groups. (It should be noted The diversity of Oligocene and Miocene that there has been massive extinction of an- fossils for the subfamily Rhyparochrominae giosperms in Africa [Raven & Axelrod, (Scudder, 1890; Statz and Wagner, 1950) and 1974]). But, when Afrovertanus is viewed in their frequent similarity to modern forms, the complete context of the tribal phylogeny, strongly suggests that this seed-feeding in- it seems more likely that its occurrence in sect group arose in the Mesozoic and radiat- Africa is accounted for by the secondary col- ed during the Cretaceous coincident with the onization mentioned above. radiation of the angiosperms. Preliminary When only the Western Hemisphere is but convincing sister group evidence for liv- considered, the distributions of Kolenetrus, ing forms does suggest Gondwanaland the Nearctic monophyletic group including origins for some of the more primitive tribes Prytanes, Type III and the non-pruinose ele- (Slater and Sweet, 1970; Sweet, unpubl.; ment of Type IV (from hypothetical ancestor Harrington, unpubl.). 37) all attest to a long period of isolation of The Myodochini, a highly derived tribe North America. Rosen's (1975) vicariance within the subfamily, which Sweet (1967) has model of Caribbean biogeography, while fo- postulated as arising "from a southern hemi- cusing on the Antillean fauna, does summa- sphere Udeocorini ancestry,." probably orig- rize Malfait and Dinkleman's (1972) recent inated in west Gondwanaland in the Late plate tectonic evidence of a Late Jurassic 1980 HARRINGTON: MYODOCHINI 67

archipelago connecting the Americas which myodochine stocks mentioned above did was subsequently displaced to form the An- reach North America and were able to tilles. This connection, and indeed the early evolve there in comparative isolation before phases of its eastward displacement, would the Pliocene closure of the Isthmus of Pan- have afforded potential avenues for faunal ama, while their South American ancestors exchange during a prolonged period in the in each case were probably supplanted by Early Tertiary. Whatever the avenue or time the radiation of the more dominant Neotrop- of entry, the ancestors of each of the four ical myodochine lineages extant today.

CLASSIFICATION Slater's (1964a) catalogue included 40 gen- includes Afrovertanus Scudder as a new my- era in the tribe Myodochini. From these 40 odochine genus. Sweet (1967) removed Caridops to the Rhy- Since Slater (1964a), Cholula has been parochromini, Insulicola to the Udeocorini moved into the Myodochini by Sweet (1967) and Phaeax to the Bledionotinae, and he has and Fontathanus Scudder (1963), Tenuicoris recently (1977) placed Neosuris in the Udeo- Slater and Harrington (1974) and Megapa- corini. mera Scudder (1975) have been described. Sweet (1967) retained Altomarus in the Recently, Malipatil (1978) has revised the Myodochini, stressing that ".... Although tribe Myodochini for the Australian region having spiracle 2 ventral, . . . It lacks inner and described the new genera Henicorthaea, laterotergites, has a round pronotum, and an Horridipamera, Pamerapa, Pamerarma, abdominal type stridulitrum." To retain Al- Paraeucosmetus, and Pseudopachybrachius tomarus in the Myodochini, however, one which are included in this revision and cla- must argue a secondary ventralization of spi- distic analysis. In the same paper he also racle 2. As the dorsal abdominal sclerotiza- describes three other genera, Exopamera, tion of this genus is very light and apparently Myodorthaea, and Woodwardocoris on re- reduced, one can easily hypothesize the loss spectively, only one, two, and four female of inner laterotergites (they are often quite specimens. In light of the importance of male faint in other taxa). I prefer this second hy- genitalia in the classification of the Myodo- pothesis and place Altomarus in the tribe chini, I have not included these last three where it appears to be re- genera, based on such limited material, in my lated to Caridops with which it shares a high- study. Malipatil (1978) has also synonymized ly shining body surface, an abdominal stri- Remaudiereana as a junior synonym of dulitrum and peculiarly cleft humeral angles Pachybrachius. I do not believe that this ac- not seen among the Myodochini. tion is at all warranted (see generic diagnoses No modern workers have examined Fon- for discussion) and thus am elevating Re- tejanus wasmanni Breddin (1904), and I was maudiereana to generic status once again. unable to locate the type specimen. Com- Additionally, in this study Sphaerobius is parison of Distant's (1910) figure of the type considered to be a junior synonym of Erlac- and habitat notes from the original descrip- da, Neocattarus a junior synonym of Chol- tion with the figure and habitat description ula, Exptochiomera ajunior synonym ofPry- of the udeocorine genus Serranegra Lind- tanes, and Orthaea is elevated to generic berg strongly suggests that they are synon- status again. Thirteen new genera are rec- ymous. I consider Fontejanus a udeocorine ognized (12 for previously described species). and excluded it from this analysis. The following key includes 56 myodochine The addenda to Slater's (1964a) catalogue genera. 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

FIGS. 2-8. 2. Heraeus plebejus, dorsal view. How to measure: a. Width across eyes. b. Interocular distance. c. Distance between ocelli. d. Width anterior pronotal collar. e. Width transverse impression. f. Postocular distance. g. Distance from base of head to antennal insertion. h. Length of head (nonde- clivent). i. Length anterior pronotal lobe (including collar). j. Length posterior pronotal lobe. k. Total length. 3. Kolenetrus plenus, lateral view of head. 1. How to measure length of a declivent head. 4. "Sharp" pygophore, schematic. 5. "Subsharp" pygophore, schematic. 6. "Broadly rounded pygophore, schematic. 7. Pygophore "with median depression in posterior lip," schematic. 8. Pygophore "with distinct median groove in posterior lip," schematic. 1980 HARRINGTON: MYODOCHINI 69

TRIBE MYODOCHINI STAL propleuron; if brachypterous, mem- Generally with pronotum rounded brane not shaped as above ...... 6 laterally 6(5). Ocelli absent ...... Cnemodus (fig. 98) and distinctly bilobed; sterna IV and V Ocelli present ...... 7 fused, with the intersegmental suture usually 7(6). A crescent-shaped, striated area present curving cephalad laterally; abdominal spira- ventrolaterally on anterior portion of cles II through IV dorsal, the remainder ven- abdomen ...... 8 tral; inner laterotergites absent (present in Striated area lacking on anterior portion the Udeocorini, the only other rhyparo- of abdomen ...... 14 chromine tribe with spiracles II through IV 8(7). Head prolonged with a distinct "neck" dorsal); nymphs with three dorsal abdominal region; striated area confined to ab- scent glands and abdomen not heavily scler- dominal sterna II and III (fig. 28); plec- otized; Y-suture well in trum on hind femur consisting of two developed nymphs. chisel-like projections (fig. 29)......

...... Erlacda (fig. 85) KEY TO THE GENERA OF THE Head not prolonged into a neck; striated TRIBE MYODOCHINI1 area extending onto sternum IV (fig. 1. Interocular distance less than postocular 35); plectrum on hind femur a line or distance ...... 2 scattered field of minute tubercles (fig. Interocular distance equal to or exceeding 36) ...... 9 postocular distance ...... 5 9(8). Three rows of claval punctures (fig. 19); 2(1). Entire pronotum pruinose in dorsal view; anterior pronotal lobe shining except (New World) ...... 3 collar, collar and posterior lobe prui- At least anterior lobe of pronotum shining nose; stridulitrum obscure, difficult to in dorsal view; (Old World) ...... 4 detect, extending only onto anterior 3(2). Posterior margins of ocelli located ante- portion of sternum IV ...... rior to posterior margins of eyes; eyes ...... Slaterobius(fig. 69) longitudinally oval ...... More than three rows of claval punctures (figs. 17, 18); entire protonum uniform- ...... Pephysena (fig. 86) of ocelli located ly pruinose, shagreened or shining; stri- Posterior margins poste- dulitrum either conspicuous or obscure rior to posterior margins of eyes; eyes but basically round ... Myodocha (fig. 97) always extending well onto sternum curved IV (fig. 35) ...... 10 4(2). Scutellar apex upward, bluntly 10(9). Length antennal segment I greater than thornlike; claval punctation in three interocular regular rows (fig. 19); lacking a cres- distance; pronotum shining striated area or shagreened, not pruinose and never cent-shaped laterally on deeply punctate; male fore tibia typi- abdominal sterna II and III ...... cally adorned with a spine or spines ...... Aegyptocoris (fig. 43) ...... 11 Scutellum uniformly flat, not elevated api- Length antennal more than four rows of claval segment I less than in- cally; terocular distance; pronotum typically punctures (fig. 17); a crescent-shaped, pruinose, in some cases very deeply dark, striated area present laterally on punctate; male fore abdominal sterna II and III (first two tibia unarmed ...... 12 visible sterna) (fig. 33)...... 11(10). Body subshining, shagreened; hemelytra ...... Afrovertanus (fig. 88) 5(1). A distinctive cross-striated area present often coleopteroid, stridulitrum dis- on propleuron, curving anteroventrad cerned with difficulty ...... from pleural suture to midventral an- ...... Ashlockaria (fig. 90) terior margin of collar (fig. 39); often Body highly shining; hemelytra usually brachypterous with macropterous; stridulitrum conspicu- characteristically ous, shining, often slightly raised and shaped membrane (fig. 30) ...... darker than rest of abdomen ...... Pseudocnemodus (fig. 84) ...... No curving cross-striated area present on ...... Pseudopamera(fig. 94) 12(10). Stridulitrum diffuse, not readily apparent; 'Figures 2 and 3 indicate how to make the various pronotum and head deeply punctate; measurements indicated in this key. abdomen with a band of long silvery 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

\\ .13 15

16 FIGS. 9-16. 9. Type I phallus, schematic. 10. Type II phallus, schematic. 11. Type III phallus, schematic. 12. Type IV phallus, schematic. 13. Fore femur multispined. 14. Fore femur with pronounced median spine. 15. Fore femur with spines double-ranked. 16. Fore femur with spines only along inner edge.

hairs covering much of sternum IV in deeply incised; always macropterous;

lateral view .... Stridulocoris (fig. 59) antennal segment IV with a light prox- Stridulitrum fairly readily apparent; imal band; fore femoral spines clearly pronotum and head not deeply punc- double ranked (spines present on both tate; abdomen of uniform vestiture with inner and outer edge of ventral surface) no area of long silvery hairs ...... 13 (fig. 15) ... Froeschneria (fig. 67) 13(12). Lateral margin of posterior pronotal lobe Lateral margin of posterior pronotal lobe angled posterolaterad at approximately angled posterolaterad at less than a 45 a 45 degree angle; pronotum markedly degree angle; transverse impression not bilobed with transverse impression deeply incised; often submacropterous; 1980 HARRINGTON: MYODOCHINI 71

V. .~~~ Yb.x7* X 9 * .'2

17

23

25

26

~~1Z2 ~~~~2

FIGS. 17-36. 17. Four or more rows of claval punctures. 18. Three regular rows and a few additional punctures. 19. Three regular rows of claval punctures. 20. Membranal edge of corium impunctate. 21. Membranal edge of corium with a row of punctures. 22. Lateral pronotal carina, Suffenusfusconervosus. 23. Clavate antenna, Carpilis barberi. 24. Capitate antenna, Ptochiomera nodosa. 25. Capitate antenna, Sisamnes clavigera. 26. Scalloping prolonged as cellular division of sternum II. 27. Edge of abdominal sternum II finely scalloped. 28. Stridulitrum, Erlacda arhaphaeoides. 29. Plectrum, Erlacda arhap- haeoides. 30. Membrane shape, Pseudocnemodus canadensis. 31. Stridulitrum, Stridulocoris gracilis. 32. Plectrum, Stridulocoris gracilis. 33. Stridulitrum, Afrovertanus elongatus. 34. Plectrum, Afrover- tanus elongatus. 35. Stridulitrum, Ligyrocoris sylvestris. 36. Plectrum, Ligyrocoris sylvestris. 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

0~~~~~

49 ~ ~~ - 50

X~47~~~~~~~~~~~~~~~~~~~~~~5A0t'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

52

FIGS. 37-54. 37. Mesepimeron "enclosed." 38. Mesepimeron "emergent." 39. Stridulitrum, Pseu- docnemodus canadensis. 40. Plectrum, Pseudocnemodus canadensis. 41. Evaporative area "reduced." 42. Evaporative area "extensive." 43. Aegyptocoris coatoni, dorsal view. 44. Pamerapa thoracicus, dorsal view. 45. , dorsal view. 46. Fontathanus flavonotus, dorsal view. 47. Stalaria ferruginosus, dorsal view. 48. Sisamnes clavigera, dorsal view. 49. Prytanes minima, dorsal view. 50. Cholula bicolor, dorsal view. 51. Caenopamera forreri, dorsal view. 52. Remaudiereana tibialis, dorsal view. 53. Pamerana nigritula, dorsal view. 54. Henicorthaea yeoi, dorsal view. 1980 HARRINGTON: MYODOCHINI 73

antennal segment IV usually uniformly 18(17). Pronotum dorsoventrally compressed to dark, if light banded, then fore femoral form a lateral carina, this most apparent spines single ranked (present only along in area of transverse impression but inner edge of ventral surface of femur) clearly extending onto both lobes; an- (fig. 16) ...... Ligyrocoris (fig. 68) tennal segment III slender, longer than 14(7). Evaporative area reduced (fig. 41); claval interocular distance ......

punctation usually in three rows (fig...... Ereminellus(fig. 65) 19); pronotum usually shining, never Pronotum typically ecarinate, if subcari- pruinose; in most cases length of anten- nate, only bluntly so on posterior lobe; nal segment III less than interocular length antennal segment III usually less distance ...... 15 than interocular distance, if greater, Evaporative area not reduced as above; then also extremely swollen ...... 19 claval punctation in more than three 19(18). Antennal segment III markedly enlarged rows (figs. 17, 18); pronotum dull, sha- and swollen, distally of greater diame- greened or pruinose; antennal segment ter than segment I; lateral margin of III usually longer than interocular dis- posterior pronotal lobe bluntly subcar- tance ...... 22 inate ...... 20 15(14). No collar apparent on anterior pronotal Antennal segment III generally filiform, lobe; ventral surface of head not if somewhat clavate, of no greater di- grooved; buccular juncture U-shaped, ameter distally and often more slender but not near labial insertion, occurring than segment I; lateral margins of at level of eyes (fig. 79) ...... pronotum ecarinate ...... 21 ...... Kolenetrus (fig. 58) 20(19). Anterior pronotal lobe globose, strongly An anterior pronotal collar apparent but convex in lateral view; transverse not demarked posteriorly; ventral sur- impression complete and distinct; an- face of head grooved; buccularjuncture tennal segments II, III and IV approx- U-shaped close to labial insertion (fig. imately equal in length ......

80) ...... 16 ...... Ptochiomera (fig. 56) 16(15). Antennal segment II very elongate, great- Anterior pronotal lobe with lateral mar- er in length than segments III and IV gins rounded but not globose, nearly combined, also longer than twice inter- flat when viewed laterally; transverse ocular distance; only collar of anterior impression obsolete mesally; antennal pronotal lobe punctate ...... segment IV distinctly longer than either ...... Caenopamera (fig. 51) II or III ...... Sisamnes (fig. 48) Length antennal segment II less than 21(19). Both eyes and ocelli very rounded and combined lengths of segments III and beadlike, markedly elevated from head IV and less than twice interocular dis- surface; length antennal segment II less tance; entire anterior pronotal lobe than interocular distance ......

punctate (punctures may be smaller and ...... Valonetus(fig. 61) less apparent than those of posterior Eyes and ocelli not abnormally elevated pronotal lobe) ...... 17 from head; length antennal segment II 17(16). All antennal segments generally stout, greater than interocular distance .....

bearing characteristic long hairs per- ...... Prytanes(fig. 49) pendicular to long axis of antennae; 22(14). Pronotum dorsoventrally compressed with wings uniformly pale with small evenly a distinct lateral carina on both lobes . . distributed punctures darker than but ...... 23 not strongly contrasting with back- Lateral pronotal margins ecarinate . . . 24 ground of clavus and corium ...... 23(22). Extremely small, approximately 2.5 mm...... Carpilis (fig. 57) long or less; claval punctures in three Antennae usually filiform or capitate with regular rows (fig. 19); scutellum uni- segments III and IV swollen, bearing formly colored and flat; (Old World).. only fine ...... Suffenus(fig. 55) apically directed hairs; wings Large, considerably more than 2.5 mm. with dark pigmentation marking specif- long; more than three regular rows of ic patterns against lighter background claval punctures (fig. 17); scutellum and large dark punctures irregularly ar- with a low median carina, apex pale; ranged with some areas impunctate ... (South America) ......

...... 18 ...... Megacholula(fig. 62) 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

24(22). Both lobes of pronotum and head uni- Mesepimeron enclosed by metepisternum formly and coarsely punctate; head touching mesepisternum (fig. 37) .. 36 broad, appearing almost to rest with 29(28). Head elongate, often with a distinct eyes touching anterior pronotal angles; "neck" portion; postocular distance sternal scalloping prolonged in a cellu- equal to or greater than distance be- lar pattern, clearly visible on sternum tween ocelli ...... 30 II (fig. 26); a very narrow anterior pro- Head less elongate, never with a distinct notal collar vaguely indicated but never "neck" portion; postocular distance demarked posteriorly by a linelike less than distance between ocelli .. 34 groove ...... 25 30(29). Very large (ca. 20.0 mm. long); labium Typically with head and anterior pronotal reaching well onto abdomen; fore tibia lobe impunctate or only vaguely punc- armed with several small spines in both tate; head generally with eyes removed sexes ...... Megapamera (fig. 82) from anterior pronotal angles; sternal Much smaller (less than 10.0 mm.); la- scalloping usually lacking, if present, bium not attaining abdomen; fore tibia largely hidden under metapleuron and unarmed ...... 31 not prolonged; in most cases a distinct 31(30). Eyes longitudinally oval; posterior margin anterior pronotal collar, demarked pos- of ocelli located distinctly anterior to teriorly by a linelike groove present.. posterior margin of eyes ...... 32

...... 27 Eyes rounded; posterior margin of ocelli 25(24). Claval punctation in three regular rows clearly located posterior to posterior (fig. 19); apex of scutellum rounded, margin of eyes ...... 33 elevated and abruptly pale; bucculae 32(31). Head with a clearly defined cylindrical

with a posterior projection (fig. 78) ... "neck"; vertex of head convex; ante-

...... Paracholula (fig. 64) rior pronotal lobe pruinose ...... Often with more than three rows of claval ...... Pephysena(fig. 86) punctures; scutellum unicolorous; buc- "Neck" region of head less definitely de- culae not projecting posteriorly (fig. fined; vertex of head flat, depressed be- 26 79) ...... tween eyes; anterior pronotal lobe shin- 26(25). Abdomen with a large glabrous area on ing ...... Tenuicoris (fig. 96) lateral portions of sterna II and III 33(31). Ocellus either approximately equidistant (microtexture filelike, this a diffuse stri- from posterior margin of head and eye dulitrum); hind femur with a plectrum or closer to posterior margin of head; of scattered spines on basal one-half; head essentially flat when viewed lat- abdomen often with a band of long sil- erally, not showing a gradual rounded very hairs covering much of sternum IV constriction from eyes to insertion of in lateral view; lateral pronotal margins head; fore coxae unspined; ventral por- ecarinate ...... Stridulocoris (fig. 59) tion of collar not produced anteriorly

Abdomen with uniform vestiture, no shin- ...... Catenes(fig. 93) ing stridulitrum or area of long hairs as Distance from posterior margin of head to above; hind femur devoid of spines; ocellus greater than (often two or three posterior pronotal lobe with a blunt ca- times) distance from ocellus to margin rina, sometimes forked over humeral of eye; head when viewed laterally angles ...... Cholula (fig. 50) showing gradual rounded constriction 27(24). A distinctive row of corial punctures from eyes to insertion of head; fore along membranal margin (fig. 21); fore coxae armed with a spine; ventral por- leg very slender; fore coxal spine lack- tion of collar produced anteriorly ...... ing; fore femoral spines present only ...... Heraeus (fig 87)

along inner edge of ventral surface ... 34(29). Jugum rounded; length of head greater

...... Stigmatonotum (fig. 63) than or equal to 2.5 times interocular Lacking a membranal margin row of cor- distance; head elongate with large eyes; ial punctures (fig. 20); usually with a (Old World) ...... Pamerana (fig. 53) fore coxal spine; spines ranked along Jugum forming a fine ridge above antennal both inner and outer edge of ventral segment I; length of head less than 2.5 28 surface of fore femur ...... times interocular distance; head some- 28(27). Mesacetabulum with mesepimeron emer- times elongate, but not as in figure 53; gent from between meso- and metepi- (New World) ...... 35 sternum (fig. 38) ...... 29 35(34). Head and anterior pronotal lobe lower 1980 HARRINGTON: MYODOCHINI 75

than posterior pronotal lobe; fore coxal notal lobe and head also often punctate, spine poorly developed or absent; color though sometimes with punctures mi- predominantly blackish brown with nute and not readily discernible .. . 46 characteristic orange areas along the 41(40). Pronotal transverse impression shallow, lateral margins of posterior pronotal obsolete medially; (New World) ...... lobe and paired orange maculae medi- ...... Perigenes (fig. 72) ally on the posterior pronotal lobe ad- Transverse impression complete, narrow, jacent to the transverse impression ... linelike, deeply incised; (Old World)..

...... Orthaea (fig. 91) ...... 42 Dorsal surface of head and both lobes of 42(41). Total length less than 3.5 times width of pronotum in essentially same plane; posterior pronotal lobe ...... 43 fore coxal spine(s) well developed; Total length greater than 3.5 times width lacking the characteristic orange mark- of posterior pronotal lobe ...... 44 ings described above ...... 43(42). Pronotum evenly pruinose but devoid of ...... Neopamera (fig. 83) long hairs; fore femur markedly incras- 36(28). Head very broad, interocular distance sate; a small spine midlength on the equal to or greater than width of pro- male fore tibia; labial segment I attain- notal collar ...... 37 ing base of head .. Pamerapa (fig. 44) Head not especially broad, interocular Body with many very evident long hairs distance less than width of pronotal col- in addition to pruinosity; fore femur rel- lar ...... 38 atively slender; male fore tibia un- 37(36). Strikingly ant mimetic with lateral margin armed; labial segment I clearly not at- of head between eye and insertion of taining base of head......

antenna expanded as a platelike curving ...... Humilocoris (fig. 60) ridge; a rounded but distinct spine pres- 44(42). Posterior margin of pronotum straight ent on lateral margin of pronotal col- across base of scutellum; male fore tib- lar ...... Xenydrium (fig. 92) ia with at most two tiny spines near dis- Lateral margin of head between eye and tal end; collar with at least a few faint base of antenna not expanded in a punctures; (Palearctic) ......

ridge; pronotal collar unspined ...... Pachybrachius (fig. 45) ...... Distingphyses (fig. 89) Posterior margin of pronotum concave 38(36). Head with essentially no postocular re- across base of scutellum; male fore tib- gion, eyes seeming to touch anterior ia curving with a single spine near mid- pronotal angles; no clearly demarked dle (often tibia strongly curving and anterior pronotal collar apparent ..... spine large); collar impunctate, ring- ...... Fontathanus (fig. 46) like; (Old World tropics) ...... 45 Head not extremely elongate but with 45(44). Body robust, subovoid with anterior pro- eyes clearly removed from anterior notal lobe globose; width of head pronotal angles and with a postocular across eyes less than width of trans- region apparent; anterior pronotal lobe verse impression; male fore tibia typi- with a very distinct anterior collar de- cally curving and with a large spine marked posteriorly by a linelike midlength; male genitalia with holding groove .... 39 sclerites .... Remaudiereana (fig. 52) 39(38). Large (ca. 9.5 mm. or longer); uniformly Body slender, elongate with anterior pro- dark, even hemelytra unpatterned; en- notal lobe less globose; width of head tire body surface dull, shagreened, not across eyes equal to or often greater pruinose . Zeropamera (fig. 95) than width of transverse impression; Body and hemelytra usually showing col- male fore tibia relatively straight with or patterning, sometimes totally pale a slender spine midlength; male geni- but never uniformly dark; thorax above talia lacking holding sclerites...... and below primarily pruinose, not sha- ...... Pamerarma (fig. 99) greened .... 40 46(40). Fore femoral spines single ranked, pres- 40(39). Head and anterior pronotal lobe including ent only on inner edge of ventral sur- collar impunctate, or collar only with face (fig. 16) ...... 47 a few faint punctures; pronotal collar Fore femoral spines double ranked, pres- narrow and ringlike ...... 41 ent on both inner and outer edges of Anterior pronotal collar usually broad and ventral surface (fig. 15) ...... 48 distinctly punctate; rest of anterior pro- 47(46). Stout dark bristles present on all tibiae; 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

fore coxal spine distinct; first segment 52(51). Small (ca. 4.0 mm. or less) and ant mi- of hind tarsus approximately 3 times metic; head strongly swollen in lateral the combined lengths of segments 2 and view; eyes small, rounded, protruding 3; (New World) ...... and beadlike ...... 53

...... Zeridoneus (fig. 70) Typically larger insects 5.0 mm. or great- Fore tibia lacking such stout dark bristles; er; if small, then not ant mimetic with fore coxal spine absent or barely indi- a swollen head and beadlike eyes.. 54 cated; first segment of hind tarsus ap- 53(52). Hemelytra predominately dark with light proximately 2 times the combined patterning; anterior two thirds of ante- lengths of segments 2 and 3; (Old rior pronotal lobe across calli shining; World) ...... Stalaria (fig. 47) (Old World) ...... Mimobius (fig. 75) 48(46). Anterior pronotal lobe, including collar, Hemelytra predominately pale with dark shorter than posterior lobe; fore femo- patterning; anterior pronotal lobe prui- ral spines short, stout and pale with nose, not shining across calli; (New

dark apices ... Henicorthaea (fig. 54) World) ...... Bacacephalus (fig. 102) Anterior pronotal lobe, including collar, 54(52). Transverse pronotal impression wider longer than posterior lobe; fore femoral than head; head as long as or longer spines relatively long, slender and uni- than width across eyes; juga rounded

...... Para aromius fi . 73 colorous ...... 49 ...p ( g ) 49(48). Head elongate with large eyes (fig. 53); Head wider than long and as wide as or width of head across eyes more than wider than transverse impression; a twice interocular distance; pygophore minor jugal ridge usually distinguish- with a deep median longitudinal groove able ...... 55 in the posterior lip (fig. 8); phallus of 55(54). Anterior pronotal lobe globose, impunc-

Type I (fig. 9) .... Pamerana (fig. 53) tate, shagreened; distance base of head Head usually not noticeably elongate; to insertion of antenna exceeding inter- width of head across eyes less than ocular distance; transverse impression twice interocular distance; posterior and posterior demarcation of collar edge of pygophore broadly rounded deeply incised ......

. . (fig. 6), lacking a median groove; phal- ...... Horridipamera (fig 74) lus of Type II (fig. 10) ...... 50 Anterior pronotal lobe punctate and gen- 50(49). Pronotum tapering cephalad with anterior erally not extremely globose; distance lobe flattened and very little convex, in base of head to insertion of antenna lateral view anterior lobe distinctly generally less than the interocular dis- lower than posterior lobe; collar with tance (if more, anterior pronotal lobe characteristic median "dip" to poste- always at least faintly punctate); trans- rior margin (fig. 66); abdomen equal to verse impression and posterior demar- or longer than combined length of head cation of collar not very deeply in- and pronotum ..... Paromius (fig. 66) cised ...... 56 Anterior pronotal lobe usually strongly 56(55). Large, usually greater than 6 mm. in convex, in lateral view not lower than length; head broad and jugal ridge posterior lobe; collar essentially straight above antennal segment I distinct . 57 across midline, not as in figure 66; ab- Small, generally less than 5 mm. in length; domen shorter than combined length of jugal ridge above antennal segment I head and pronotum ...... 51 usually very narrow and poorly devel- 51(50). Generally brachypterous; length anterior oped .. Pseudopachybrachius (fig. 77) pronotal lobe equal to or greater than 57(56). Anterior pronotal lobe globose and im- 2.5 times length posterior pronotal punctate; male fore tibia spined ...... lobe; male fore tibia curved and bearing ...... Eucosmetus (fig 71) one prominent spine and other minor Anterior pronotal lobe punctate in dorsal spines on distal one-half ...... view at least laterally and toward trans-

...... Togo (fig. 76) verse impression; male fore tibia not Generally macropterous; male fore tibia spined ...... 58 not noticeably curved and often un- 58(57). Usually with a broad dark band across armed, if armed, then length anterior hemelytra at level of corial apex and a pronotal lobe less than 2.5 times length second more anterior dark area present posterior lobe ...... 52 and reaching the lateral corial margin 1980 HARRINGTON: MYODOCHINI 77

at level of claval apex; male genitalia second more anterior dark area either with apical conjunctival spines sym- lacking or present only medially and metrical, an additional pair of more not reaching lateral corial margin at proximal pigmented or sclerotized level of claval apex; male genitalia with lobes present on conjunctiva; (Old left apical conjunctival spine bifid, no World) ... Paraeucosmetus (fig. 100) additional spines or lobes on conjunc- A broad dark band extending across hem- tiva; (New World) ......

elytra at level of corial apex but any ...... Pseudoparomius(fig. 101)

GENERIC DIAGNOSES AND DESCRIPTIONS The following generic diagnoses and brief odochine genus to show no indication of an summary descriptions are presented in the anterior pronotal collar in dorsal view. It is order in which the genera appear in the also plesiomorphic for most other characters cladogram. After each name in the species considered in this study. lists my familiarity with that species is indi- Apomorphic states further characterizing cated by the following system: the genus include the highly shining body surface, reduced evaporative area and re- * I saw determined material. duction of the fore femoral spines. Although ** I saw the holotype or lectotype (in a few these apomorphic character states (all of in- cases saw only a paratype). dependent derivation) and its general habitus Generally, I saw no material for the and size might lead one to associate Kole- species and thus am uncertain of generic netrus with the monophyletic Nearctic group placement. with a Type I phallus, it is readily distin- The original reference for each species and guished from all those genera by its lack of all subsequent literature, including synony- a collar, buccular condition, and lack of a mies, that are cited in Slater's 1964a cata- midventral groove on the head. logue are not repeated in this paper. How- DESCRIPTION: Body small, ovoid, heavi- ever, new species (most to be described ly punctate and highly shining; pronotum elsewhere), new combinations, new synon- dorsoventrally compressed, lateral margins ymies, general geographic distribution of subcarinate; no anterior collar apparent dor- each genus and any nomenclatural changes sally. subsequent to Slater's 1964a catalogue are Phallic Type I (fig. 9); posterior edge of all indicated for each genus. pygophore sharp (fig. 4); claval punctation in three regular rows (fig. 19); ventral surface of head not grooved; buccular juncture broadly U-shaped occurring close to base of KOLENETRUS BARBER head and enclosing a gular area (fig. 79); Figure 58 evaporative area reduced (fig. 41); mesepi- TYPE SPECIES: plenus meron enclosed (fig. 37); anterior margin of Distant, 1882.** abdominal sternum II (first sternum visible) Monotypic, broadly distributed in Nearc- scalloped (fig. 27); fore femur with spines tic, south to Guatemala. only along inner edge of ventral surface (fig. DIAGNOSIS: Kolenetrus is the only my- 16). 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

58

62

FIGS. 55-65. 55. Suffenus fusconervosus, dorsal view. 56. Ptochiomera nodosa, dorsal view. 57. Carpilis barberi, dorsal view. 58. Kolenetrus plenus, dorsal view. 59. Stridulocoris gracilis, dorsal view. 60. Humilocoris cephalotes, dorsal view. 61. Valonetus pilosus, dorsal view. 62. Megacholula engle- mani, dorsal view. 63. Stigmatonotum sparsum, dorsal view. 64. Paracholula vegetus, dorsal view. 65. Ereminellus fusicornis, dorsal view.

MEGACHOLULA, NEW GENUS DIAGNOSIS: Megacholula, Suffenus, and Figure 62 Ereminellus are the only three myodochine genera that show a distinct continuous carina TYPE SPECIES: Megacholula englemani, along the lateral margins of both pronotal new species. lobes. Size immediately distinguishes Me- Monotypic, Neotropical. gacholula (total length greater than 7 mm.) 1980 HARRINGTON: MYODOCHINI 79 from the other two carinate genera which are gin; posterior pronotal lobe, scutellum and approximately half that length. Suffenus and hemelytra pruinose, light tawny with chest- Ereminellus also both have the claval punc- nut punctures; membrane of hemelytra hya- tation in three regular rows (fig. 19), whereas line; lateral carina on posterior pronotal lobe Megacholula shows four irregular rows of pale buffy yellow; antennae light buffy claval punctures (fig. 17). brown, gradually darkening to between buffy DESCRIPTION: Head shining, impunctate brown and chestnut on segment IV; legs and and acuminate, broad across eyes, interoc- labium light yellowish buffy brown, numer- ular area flattened. Pronotum declivent from ous small sepia spots of heavier sclerotiza- posterior margin, dorsoventrally compressed tion on femora; thorax ventrally and laterally with a distinct carina along the entire lateral light chestnut. margin of both pronotal lobes; shape of Length of head .87 mm., width across eyes pronotum subrectangular, anterior angles 1.45 mm., interocular distance .84 mm., nearly right angles, giving a very square- length anterior pronotal lobe .95 mm., length shouldered appearance; lateral margins little posterior pronotal lobe .84 mm., maximum sinuate and nearly parallel; transverse width of anterior pronotal lobe 1.63 mm., impression barely apparent; an anterior pro- width of transverse impression 1.56 mm., notal collar detectable only medially; poste- width across humeral angles 1.98 mm.; rior margin slightly convex across base of length of scutellum 1.18 mm., width of scu- scutellum; pruinose anterior lobe impunc- tellum 1.14 mm.; length of corium 3.23 mm., tate, posterior lobe with numerous moderate distance apex corium to apex membrane 1.22 sized punctures and pruinose. Scutellum mm.; labium very long, reaching base of ab- with a thin, pale, impunctate median longi- dominal sternum III, length labial segments: tudinal stripe and paler swollen apex; re- I 1.10 mm., 11 1.22 mm., III .84 mm., IV .95 mainder of scutellum punctured like poste- mm.; length antennal segments: I .53 mm., rior pronotal lobe. Hemelytra slightly 11 1.33 mm., III 1.14 mm., IV 1.22 mm.; total exceeding abdominal apex; clavus with length 7.33 mm. punctures in four irregular rows (fig. 17); lat- HOLOTYPE: Ecuador: d Limoncocha, eral corial margins essentially straight. Legs March 1974, (H. Dodge Engleman), in the elongate; fore coxa with two small spines; American Museum of Natural History. fore femur incrassate, multispined as in fig- PARATYPES: Y Same data as above, in J. ure 13; fore tibia unarmed in both sexes; A. Slater collection. d Same data as above, mesepimeron enclosed (fig. 37); evaporative in J. Harrington collection. area extensive (fig. 42); buccular juncture ETYMOLOGY: This new species is named broadly U-shaped, occurring at level of pos- for Dr. H. Dodge Engleman of the Coco Solo terior margin of eye and enclosing a gular Hospital, Canal Zone. Dr. Engleman has area (fig. 79). Antennae elongate, filiform. been diligent in collecting and has generously Male genitalia of Type I (fig. 9); posterior sent me a rich variety of Neotropical my- edge of pygophore sharp (fig. 4). odochines, including this new species which ETYMOLOGY: From the Greek mega, bears his name. large or great, and Cholula, for the appear- ance of this genus which is superficially like PARACHOLULA, NEW GENUS a giant specimen of the genus Cholula. Figure 64 TYPE SPECIES: Neocattarus thoracicus Megacholula englemani, new species Distant, 1882. DIAGNOSIS: This is the only described Two species, Neotropical. species in the genus. INCLUDED SPECIES: [Both species are DESCRIPTION: Head and anterior pro- new combinations (from Neocattarus).] notal lobe tawny brownish red, impunctate; thoracicus (Neocattarus) Distant, head shining; anterior pronotal lobe sha- 1882.** greened, darker toward carinate lateral mar- vegetus (Neocattarus) Distant, 1882.** 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

FIGS. 66-77. 66. Paromius gracilis, dorsal view. 67. Froeschneria multispinus, dorsal view. 68. Ligyrocoris sylvestris, dorsal view. 69. Slaterobius insignis, dorsal view. 70. Zeridoneus costalis, dorsal view. 71. Eucosmetus incisus, dorsal view. 72. Perigenes dispositus, dorsal view. 73. Paraparomius leptopoides, dorsal view. 74. Horridipamera neitneri, dorsal view. 75. Mimobius sp., dorsal view. 76. Togo hemipterus, dorsal view. 77. Pseudopachybrachius vincta, dorsal view.

=pictus (Coreus) Fabricius, 1803. DIAGNOSIS: This new genus is erected to Transferred to Neocattarus from Pto- include the previously described species chiomera and synonymized by Scud- Neocattarus vegetus and thoracicus. It is der, 1970. p. 101. readily distinguished by the peculiar buccu- 1980 HARRINGTON: MYODOCHINI 81

lar condition described below and a pale, up- irrorandus (Neocattarus) Distant, turned, and swollen scutellar apex in con- 1893.** junction with the plesiomorphic state of maculatus (Neocattarus) Distant, three regular rows of claval punctures. 1893.** DESCRIPTION: Body elongate, parallel- parvus (Neocattarus) Distant, 1882.** sided; head and pronotum heavily and deep- vigens (Neocattarus) Distant, 1882.** ly punctate; pronotum only slightly com- DIAGNOSIS: Although the type species of pressed dorsoventrally, lateral margin not the genera Cholula and its junior synonym carinate; a narrow anterior collar present but Neocattarus are clearly congeneric, I have not demarked posteriorly by a linelike been unable to discover a firm synapomor- groove. phic character, other than general habitus, Phallic Type I (fig. 9); posterior edge of relating all the species included in this genus, pygophore sharp (fig. 4); claval punctation in and the group may ultimately prove not to three regular rows (fig. 19); ventral surface be holophyletic. For the present, species of of head not grooved; buccular juncture Cholula and its junior synonym Neocattarus broadly U-shaped, occurring close to base of can be recognized by the peculiar cellular head and enclosing a gular area; bucculae division of sternum II that they have in com- distinctive, well developed with a posterior mon with Paracholula and Stridulocoris but projection (fig. 78); evaporative area exten- their lack of the distinctive bucculae and scu- sive (fig. 42); mesepimeron enclosed (fig. 37); tellar apex of Paracholula and the stridula- scalloping of abdominal sternum II pro- tory mechanism found in Stridulocoris. longed as a cellular division of that segment DESCRIPTION: Body subovoid; head and (fig. 26); fore femur heavily spined with small pronotum strongly punctate; pronotum dor- spines present between rows of larger spines soventrally compressed, posterior lobe sub- occurring along both inner and outer edges carinate; anterior pronotal collar discernible of ventral surface (fig. 13). only medially and not demarked posteriorly I have not observed these insects in the by a linelike groove. field, but their general body form and col- Phallic Type I (fig. 9); posterior edge of oring suggests they may be ant or wasp mim- pygophore sharp (fig. 4); claval punctation in ics. more than three regular rows (fig. 17); ven- ETYMOLOGY: From the Greek para, be- tral surface of head not grooved; buccular side or close to, for its close relationship to juncture broadly U-shaped, occurring close the genus Cholula. to base of head and enclosing a gular area (fig. 79); evaporative area extensive (fig. 42) CHOLULA DISTANT rugose; mesepimeron enclosed (fig. 37); scal- Figure 50 loping of abdominal sternum II prolonged as Transferred to the Myodochini by Sweet, a cellular division of that segment (fig. 26); 1967, p. 224. fore femur heavily spined as in figure 13. Neocattarus Distant, 1882, NEW SYN- ONYMY. STRIDULOCORIS, NEW GENUS TYPE SPECIES: Cholula bicolor Distant, Figure 59 1882. TYPE SPECIES: Neocattarus gracilis Dis- Eight species, Neotropical. tant, 1882. INCLUDED SPECIES: bicolor Distant, Two species, Neotropical. 1882.** INCLUDED SPECIES: An undescribed discoloria Distant, 1893.** species from Mexico. variegata Distant, 1882.** gracilis (Neocattarus) Distant, 1882.** [The following five species are new com- new combination (from Neocattarus). binations (from Neocattarus).] DIAGNOSIS: The stridulatory mechanism firmus (Neocattarus) Distant, 1882.** described below has evolved independently 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

FIGS. 78-90. 78. Buccular condition Paracholula vegetus, lateroventral view of head. 79. Buccular condition Kolenetrus plenus, lateroventral view of head. 80. Buccular condition Carpilis ferruginea, lateroventral view of head. 81. Buccular condition Pephysena levis, lateroventral view of head. 82. Megapamera ashlockia, dorsal view. 83. Neopamera bilobata, dorsal view. 84. Pseudocnemodus can- adensis, dorsal view. 85. Erlacda arhaphaeoides, dorsal view. 86. Pephysena levis, dorsal view. 87. Heraeus triguttatus, dorsal view. 88. Afrovertanus elongatus, dorsal view. 89. Distingphyses insignis, dorsal view. 90. Ashlockaria sobria, dorsal view. 1980 HARRINGTON: MYODOCHINI 83

as the two species of Stridulocoris are the Phallic Type I (fig. 9); posterior edge of only stridulatory myodochines with a Type pygophore sharp (fig. 4); claval punctation in I phallus. On this basis I have given these three regular rows (fig. 19); ventral surface species generic status. Stridulocoris can also of head not grooved; buccular juncture be recognized by a characteristic band of broadly U-shaped, occurring close to base of long, dense, recumbent, silvery hairs cov- head and enclosing a gular area (fig. 79); ering much of abdominal sternum IV. evaporative area extensive (fig. 42); mesep- DESCRIPTION: Body elongate, parallel- imeron enclosed (fig. 37); anterior margin of sided; head and pronotum strongly punctate; abdominal sternum II scalloped (fig. 27); fore lateral pronotal margins rounded; a very nar- femur with spines only along inner edge of row anterior pronotal collar indicated but not ventral surface (fig. 16), these spines char- demarked posteriorly by a linelike groove. acteristically fine and sharp, closely arranged Phallic Type I (fig. 9); posterior edge of in a comblike row. pygophore sharp (fig. 4); claval punctation in more than three regular rows (fig. 17); ven- tral surface of head not grooved; buccular AEGYPTOCORIS CHINA juncture broadly U-shaped, occurring close Figure 43 to base of head and enclosing a gular area (fig. 79); evaporative area extensive (fig. 42); TYPE SPECIES: Aegyptocoris myrme- mesepimeron enclosed (fig. 37); scalloping of coides China, 1936. abdominal sternum II prolonged as a cellular Two species, Ethiopian. division of that segment (fig. 26); fore femur INCLUDED SPECIES: coatoni Slater and heavily spined as in figure 13; a diffuse ab- Sweet, 1970. p. 221.** dominal stridulitrum present (fig. 31); plec- myrmecoides China, 1936.** trum a field of minute spines on the hind fe- DIAGNOSIS: Aegyptocoris is the only my- mur (fig. 32). odochine with a Type I phallus to show head elongation, which must surely be of indepen- ETYMOLOGY: From the Latin, stridulus, dent derivation. The highly shining body and creaking or grating for this bug's sound pro- reduced evaporative area are also derived in- ducing capacity, and the Greek cori, bug. dependently in this lineage. The various characters associated with ant mimicry and SUFFENUS DISTANT the peculiar upturned scutellar apex serve to Figure 55 further distinguish this genus. TYPE SPECIES: Rhyparochromus fusco- DESCRIPTION: Body elongate, highly nervosus Motschulsky, 1863.** shining and strikingly antlike; head elongate, =Dakhla striipennis Linnavouri, 1964. prolonged behind eyes in a stalklike neck; p. 316. Synonymized by Scudder, ocelli vestigial; lateral pronotal margins 1970. p. 103. rounded; a very narrow collar area apparent Monotypic, broadly throughout Old World on anterior pronotal lobe but not demarked tropics. posteriorly by a linelike groove. DIAGNOSIS: This tiny pale monotypic ge- Phallic Type I (fig. 9); posterior edge of nus is the smallest known myodochine. In pygophore sharp (fig. 4); claval punctation in addition to its minute size it is also distin- three regular rows (fig. 19); ventral surface guished by the peculiar comblike row of of head not grooved; buccular juncture spines on the strongly incrassate fore femur. U-shaped and occurring very close to labial DESCRIPTION: Minute (ca. 2.5 mm.); insertion; evaporative area reduced (fig. 41); body ovoid; pronotum dorsoventrally com- mesepimeron enclosed (fig. 37); hemelytra pressed, lateral margins with a slight carina sometimes brachypterous; scutellum distinc- on both lobes; an anterior collar indicated tive with apex curving upwards as a large but not demarked posteriorly by a linelike blunt spine; fore coxa unspined; entire ven- groove. tral surface of fore femur heavily spined as 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

FIGS. 91-102. 91. Orthaea consuta, dorsal view. 92. Xenydrium formiciforme, dorsal view. 93. Catenes porrectus, dorsal view. 94. Pseudopamera aurivilliana, dorsal view. 95. Zeropamera nigra, 1980 HARRINGTON: MYODOCHINI 85

in figure 13; fore tibia of male straight with CAENOPAMERA BARBER many small regular teeth. Figure 51 TYPE SPECIES: Pseudopamera forreri EREMINELLUS, NEW GENUS Distant, 1893.** Figure 65 Monotypic, Nearctic. TYPE SPECIES: Exptochiomera arizonen- DIAGNOSIS: This genus is readily recog- sis Barber, 1932. nized among those genera of the Nearctic Two species, Nearctic. complex with a U-shaped buccular juncture INCLUDED SPECIES: An undescribed in a midventral head groove by its elongate species from Nevada. antennae which also bear characteristic long arizonensus Barber, 1932.** New Com- hairs perpendicular to the long axis of the bination (from Exptochiomera). antennae. Caenopamera is also larger with DIAGNOSIS: This genus shares a midven- a more elongate body form than the other tral head groove and U-shaped buccular members of that monophyletic group. juncture with the Nearctic complex involv- DESCRIPTION: Body elongate, parallel- ing Prytanes and its relatives. Yet, Eremi- sided, shagreened; lateral margins of prono- nellus is readily distinguished from members tum rounded; an anterior collar apparent but of that group by its retention of a plesio- not demarked posteriorly by a linelike morphic compressed, carinate pronotum. On groove. that basis, the former Exptochiomera arizo- Phallic Type I (fig. 9); posterior edge of nensis is recognized as a distinct genus in- pygophore subsharp (fig. 5); claval puncta- cluding an additional undescribed species tion with a few punctures in addition to three from the southwestern United States. regular rows (fig. 18); ventral surface of head DESCRIPTION: Body ovoid, shining; grooved; buccular juncture U-shaped, oc- pronotum dorsoventrally compressed, a curring within groove at level of antennal in- slight lateral carina present on both lobes; an sertion (fig. 80); mesepimeron enclosed (fig. anterior pronotal collar region clearly appar- 37); evaporative area reduced (fig. 41); fore ent but not demarked posteriorly by a line- femur with spines present only along inner like groove. edge of ventral surface (fig. 16); fore tibia of Phallic Type I (fig. 9); posterior edge of male straight with two small spines on distal pygophore subsharp (fig. 5); claval puncta- one-third; antennae extremely elongate, es- tion with a few punctures in addition to three pecially segment II which is longer than the regular rows (fig. 18); ventral surface of head combined lengths of segments III and IV. grooved; buccular juncture U-shaped, oc- curring within groove close to labial insertion (fig. 80); evaporative area reduced (fig. 41); VALONETUS BARBER mesepimeron enclosed (fig. 37); anterior Figure 61 edge of sternum II finely scalloped (fig. 27); TYPE SPECIES: Valonetus pilosus Bar- fore femur with spines present only along in- ber, 1918.** =Plociomera puberula StAl, ner edge of ventral surface (fig. 16). 1874. ETYMOLOGY: From the Greek, eremi, a Monotypic, Nearctic. desert, and the Latin, -ellus, small, for the DIAGNOSIS: The eyes and ocelli are the xeric habitat of this small, desert-dwelling most striking feature of this monotypic genus bug. of phallic Type I. They are beadlike and pro-

dorsal view. 96. Tenuicoris myrmeforme, dorsal view. 97. Myodocha serripes, dorsal view. 98. Cne- modus mavortius, dorsal view. 99. Pamerarma ventralis, dorsal view. 100. Paraeucosmetus pallicornis, dorsal view. 101. Pseudoparomius linearis, dorsal view. 102. Bacacephalus globiceps, dorsal view. 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 trude from the surface of the head, being al- close to labial insertion (fig. 80); mesepime- most substalked. The juga, which protrude ron enclosed (fig. 37); evaporative area re- like tiny tusks, are also distinctive. duced (fig. 41); anterior edge of sternum II DESCRIPTION: Body ovoid, heavily pi- finely scalloped (fig. 27); hemelytra often lose; lateral pronotal margins rounded; an brachypterous; fore femur sparsely armed anterior collar apparent but not demarked with spines only along inner edge of ventral posteriorly by a linelike groove. surface (fig. 16); male fore tibia straight and Phallic Type I (fig. 9); posterior edge of unarmed. pygophore subsharp (fig. 5); claval puncta- tion in three regular rows (fig. 19); ventral surface of head with a shallow midventral CARPILIS STAL groove; buccular juncture U-shaped occur- Figure 57 ring within groove close to labial insertion TYPE SPECIES: Carpilis ferruginea Stal, (fig. 80); bucculae relatively large, rounded 1874. and leaflike; evaporative area reduced (fig. Three species, Nearctic. 41); mesepimeron enclosed (fig. 37); anterior INCLUDED SPECIES: barberi (Ptochiom- edge of abdominal sternum II finely scal- era) Blatchley, 1924.* loped (fig. 27); hemelytra sometimes bra- consimilis Barber, 1949.** chypterous; fore femur with but a few spines ferruginea Stal, 1874.* present only along inner edge of ventral sur- DIAGNOSIS: Carpilis is readily recog- face (fig. 16). nized by its clavate antennae with all seg- ments thickened (fig. 23). Additionally, an- tennal segments II through IV have very PTOCHIOMERA SAY elongate hairs oriented at right angles to the Figure 56 long axis of the antenna. The uniformly pale, often brachypterous hemelytra, with only TYPE SPECIES: Ptochiomera nodosa Say, the punctures slightly darker than their back- 1831. ground, are also characteristic of this genus. One species, Nearctic; one species(?), DESCRIPTION: Body ovoid, shining; lat- Neotropical. eral pronotal margins rounded on both lobes; INCLUDED SPECIES: chilensis (Pachy- a distinct anterior collar present but not de- merus) Spinola, 1852.*** marked posteriorly by a linelike groove. nodosa Say, 1831.* Phallic Type I (fig. 9); posterior edge of DIAGNOSIS: Ptochiomera and Sisamnes pygophore subsharp (fig. 5); claval puncta- both have the antennae highly swollen, but tion with a few punctures in addition to three in Ptochiomera it is the third segment which regular rows (fig. 18); ventral surface of head has the greatest diameter (terminal segment with a median groove; buccular juncture most swollen in Sisamnes). The highly shin- U-shaped, occurring in groove close to labial ing body surface ofPtochiomera also serves insertion (fig. 80); mesepimeron enclosed to distinguish it from Sisamnes which has a (fig. 37); evaporative area reduced (fig. 41); peculiar granular vestiture. anterior edge of sternum II finely scalloped DESCRIPTION: Body ovoid, highly shin- (fig. 27); fore femur with spines double- ing; anterior pronotal lobe with lateral mar- ranked, present on both inner and outer edge gins rounded; posterior lobe subcarinate in of ventral surface (fig. 15); male fore tibia lateral margins; a broad anterior collar clear- curving with a well-developed spine on distal ly apparent but not demarked posteriorly by one half; hemelytra often a linelike groove. brachypterous. Phallic Type I (fig. 9); posterior edge of pygophore subsharp (fig. 5); claval puncta- SISAMNES DISTANT tion in three regular rows (fig. 19); ventral Figure 48 surface of head with a median groove; buc- TYPE SPECIES: Sisamnes contractus Dis- cularjuncture U-shaped, occurring in groove tant, 1893. 1980 HARRINGTON: MYODOCHINI 87

Two species, Nearctic; one species(?), caeca (Plociomera) Distant, 1882.** Neotropical. confusa (Exptochiomera) Barber, INCLUDED SPECIES: annulicolis (Plo- 1953.** ciomera) Berg, 1894.*** cubensis Barber, 1954b.** clavigerus (Ptochiomera) Uhler, 1895.** dissimilis (Exptochiomera) Barber, contractus Distant, 1893.** 1953.** DIAGNOSIS: Sisamnes is best character- foeda (Rhyparochromus (Plociomerus)) ized by its enlarged terminal antennal seg- Stoal, 1858.*** ments. It can be distinguished from Pto- formosa (Plociomera) Distant, 1882.** chiomera (which also has antennal segments fusicornis (Plociomera) Stal, 1874.* III and IV enlarged) by its scalelike hairs that globosus Distant, 1893.** give a granular appearance to the dorsum, intercisa (Exptochiomera) Barber, subcarinate posterior pronotal lobe, double- 1932.** ranked fore femoral spines and by antennal minima (Lygaeus (Beosus)) Guerin, segment IV being the segment of greatest 1857.* diameter. oblonga (Plociomera) Stal, 1862.* DESCRIPTION: Body ovoid, dorsum shin- plebeius (Pachymerus) Spinola, 1852. ing but given a granular appearance by many Transferred to Exptochiomera from flattened, scalelike hairs; lateral margins of Aphanus by Scudder, 1970. p. 100.*** anterior pronotal lobe rounded; lateral mar- tumens (Plociomera) Stal, 1874.** gins of posterior lobe subcarinate; a distinct DIAGNOSIS: As indicated by the dotted anterior collar present but not demarked pos- line in the cladogram (paraphyly), Prytanes teriorly by a linelike groove. is a poorly understood complex that can only Phallic Type I (fig. 9); posterior edge of be resolved by further species-level revision- pygophore subsharp (fig. 5); claval puncta- ary work. I have been unable to find a sat- tion in three regular rows (fig. 19); ventral isfactory synapomorphic condition to relate surface of head with a median groove; buc- the various species of the current genus cularjuncture U-shaped, occurring in groove Exptochiomera which is here recognized as close to labial insertion (fig. 80); mesepime- a junior synonym of Prytanes. Prytanes glo- ron enclosed (fig. 37); evaporative area re- bosus has a very globose anterior pronotal duced (fig. 41); hemelytra often brachypter- lobe (a trait commonly correlated with wing ous; anterior edge of sternum II finely reduction in the myodochines) but is other- scalloped (fig. 27); fore femur with spines wise little different from the various species double-ranked (fig. 15); male fore tibia not of Barber's genus Exptochiomera. spined; antenna capitate, the terminal two Clearly, all the species ofPrytanes belong segments strikingly enlarged with segment to the phyletic line of phallic Type I in which IV being the greatest in diameter (fig. 25). the ventral surface of the head is grooved and the U-shaped buccular juncture is close to the labial insertion. Exptochiomera = Pry- PRYTANES DISTANT tanes can only be distinguished from other Figure 49 genera in that assemblage on the basis of Exptochiomera Barber, 1928, NEW SYN- negative evidence such as the lack of anten- ONYMY. nal adaptations characteristic of Carpilis, TYPE SPECIES: Prytanes globosus Dis- Ptochiomera, and Sisamnes; the lack ofjugal tant, 1893. tusks and substalked eyes and ocelli seen in Fourteen species, Nearctic and Neotropi- Valonetus; lack of a shagreened body sur- cal. face and peculiarly haired antennae of Caen- INCLUDED SPECIES: [All species but cub- opamera and lack of the pronotal carina ex- ensis and globosus are new combinations hibited by Ereminellus. (from Exptochiomera).] DESCRIPTION: Body ovoid or subovoid, albomaculata (Plociomera) Distant, shining; lateral margins of both pronotal 1893.** lobes rounded; an anterior pronotal collar 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 apparent but not demarked posteriorly by a ly incised; lateral margins of both pronotal linelike groove. lobes rounded; a very narrow anterior collar Phallic Type I (fig. 9); posterior edge of barely apparent, not clearly demarked pos- pygophore subsharp (fig. 5); claval puncta- teriorly by a linelike groove. tion in three regular rows (fig. 19); ventral Phallic Type I (fig. 9); posterior edge of surface of head with a median groove; buc- pygophore subsharp (fig. 5); claval puncta- cularjuncture U-shaped, occurring in groove tion in four or more rows (fig. 17); buccular close to labial insertion (fig. 80); mesepime- juncture V-shaped (fig. 81); mesepimeron en- ron enclosed (fig. 37); anterior edge of ster- closed (fig. 37); evaporative area extensive num II finely scalloped (fig. 27); evaporative (fig. 42); fore femur with spines double- area reduced (fig. 41); fore femur with spines ranked (fig. 15). variable; male fore tibia with a.spine on distal one-half; hemelytra often brachypterous; an- tenna filiform. HENICORTHAEA MALIPATIL Figure 54

FONTATHANUS SCUDDER TYPE SPECIES: Henicorthaea yeoi Mali- Figure 46 patil, 1978. p. 16. Three species, Australian. TYPE SPECIES: Fontathanus elatus Scud- INCLUDED SPECIES: An undescribed der, 1963. species from New Guinea. Four species, Ethiopian. An undescribed species from the Solo- INCLUDED SPECIES: elatus Scudder, mon Islands. 1963. p. 1234.* yeoi Malipatil, 1978.*** flavonotus Scudder, 1963. p. 1234.* DIAGNOSIS: Henicorthaea is readily dis- nigronotus Scudder, 1963. p. 1235.** tinguished from other myodochines by the punctatus Scudder, 1963. p. 1235.* characteristic fore femoral spines described DIAGNOSIS: Among the Old World My- below. The peculiar relation of the head and odochini, Suffenus, Aegyptocoris, Henicor- anterior pronotal lobe to the markedly higher thaea, Humilocoris, Stigmatonotum, and posterior lobe also serves to distinguish this Fontathanus (all of phallic Type I) have the genus. anterior pronotal collar not demarked pos- DESCRIPTION: Body elongate, robust; teriorly by a linelike groove. In Suffenus the head small with large, protruding eyes; head collar area is indicated only medially. Suf- and anterior lobe of pronotum in a markedly fenus is also readily distinguished by its car- lower plane than that of posterior pronotal inate lateral pronotal margins. Aegyptocoris lobe; lateral margins of both pronotal lobes is unmistakable with its head prolonged be- rounded; a distinct anterior collar area pres- hind the eyes in a stalklike neck. ent but not demarked posteriorly by a line- Of the other four genera mentioned above, like groove. only Fontathanus has such a narrow, negli- Phallic Type I (fig. 9); posterior edge of gible collar area and the characteristic deeply pygophore sharp (fig. 4); claval punctation in bilobed pronotum the anterior angles of four or more rows (fig. 17); buccularjuncture which seem almost to touch the eyes (fig. V-shaped (fig. 81), close to labial insertion; 46). In Henicorthaea, Humilocoris, and Stig- mesepimeron enclosed (fig. 37); evaporative matonotum a fairly broad, pale collar region area extensive (fig. 42); anterior edge of ab- is readily apparent, contrasting with the rest dominal sternum II scalloped (fig. 27); fore of the dark anterior pronotal lobe. femur with spines double-ranked (fig. 15), DESCRIPTION: Body elongate, sha- spines very stout and striking, white or pale greened; pronotum elongate, especially an- in contrast to rest of fore femur and with terior lobe; transverse impression very deep- apices of spines dark. 1980 HARRINGTON: MYODOCHINI 89

HUMILOCORIS, NEW GENUS Six species: Palearctic, Ethiopian, Orien- Figure 60 tal, and Australian. INCLUDED SPECIES: afrus (Plociomera) TYPE SPECIES: Rhyparochromus cepha- Stal, 1865.* New Combination (from Pachy- lotes Dallas, 1852.** New Combination brachius). (from Stigmatonotum). capucinum (Plociomera) Stal, 1865.* Monotypic, Oriental. geniculatus (Plociomerus) Motschul- DIAGNOSIS: The Old World genera Hu- sky, 1863.* New Combination (from milocoris, Henicorthaea and Stigmatonotum Pachybrachius). (all of phallic Type I) have a distinct, broad, minutum Malipatil, 1978. p. 23.*** bandlike collar clearly indicated on the an- rufipes (Plociomerus) Motschulsky, terior pronotal lobe but not demarked pos- 1866. Transferred here by Scudder, teriorly by a linelike groove. In all three of 1970. p. 103. =japonicum (Plociom- the above genera, the collar is usually lighter era) Distant, 1883.** Synonymized by in color than the rest of the anterior pronotal Scudder, 1970. p. 103. lobe. sparsum Lindberg, 1927.* Humilocoris and Stigmatonotum lack the DIAGNOSIS: As discussed in the diagno- elaborate foreleg armature described for sis for Humilocoris, Stigmatonotum can be Henicorthaea, both having slender fore fem- distinguished from all other Eastern Hemi- ora armed with only a few spines on the inner sphere myodochines by the characteristic edge of the ventral surface. Humilocoris is row of corial punctures along the membranal slightly larger than its sister genus, Stigma- margin (fig. 21). Only Prytanes, Sisamnes, tonotum, and is distinguished from the latter Carpilis, and Ptochiomera, all Nearctic gen- by the presence of numerous long, curving, lacks era of a similar habitus to that of Stigmato- semierect body hairs. Humilocoris also notum, have similar punctation. These four the precise row of corial punctures along the that characterizes Nearctic genera have the pronotum shining membranal margin (fig. 21) dorsally, a reduced evaporative area, claval Stigmatonotum. punctation in three regular rows and a DESCRIPTION: Body subovoid; lateral U-shaped buccular juncture lying close to margins of both pronotal lobes rounded; a the labial insertion in the grooved ventral distinct anterior pronotal collar present but surface of the head. In Stigmatonotum, the not demarked posteriorly by a linelike pronotum is pruinose dorsally, the evapora- groove. edge of tive area extensive, claval punctation in four Phallic Type I (fig. 9); posterior or more rows, the head not grooved midven- pygophore broadly rounded (fig. 6); claval trally and the buccular juncture V-shaped, or more rows (fig. 17); punctation in four though close to the labial insertion. buccular juncture V-shaped (fig. 81), close to DESCRIPTION: Body elongate; lateral labial insertion; mesepimeron enclosed (fig. margins of both pronotal lobes rounded; a 37); evaporative area extensive (fig. 42); fore distinct, pale anterior pronotal collar present femur with spines only along inner edge of but not demarked posteriorly by a linelike ventral surface (fig. 16). groove. ETYMOLOGY: From the Latin humil, low Phallic Type I (fig. 9); posterior edge of or insignificant, and the Greek cori, bug, for pygophore subsharp (fig. 5); claval puncta- the undistinguished appearance of this small tion in four or more rows (fig. 17); corium brown bug. with a characteristic row of punctures pres- ent along membranal margin (fig. 21); buc- STIGMATONOTUM LINDBERG cular juncture V-shaped (fig. 81), close to la- Figure 63 bial insertion; mesepimeron enclosed (fig. TYPE SPECIES: Stigmatonotum sparsum 37); evaporative area extensive (fig. 42); fore Lindberg, 1927. femur slender with only one or a few spines 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 present on inner edge of ventral surface (fig. notatipes (Nesopamera) Barber, 1958.** 16). procera Bergroth, 1918.* punctulatus (Plociomerus) Motschul- MEGAPAMERA SCUDDER sky, 1863.* New Combination (from Figure 82 Pachybrachius). scotti (Pamera) Distant, 1901.* New TYPE SPECIES: Megapamera ashlockia Combination (from Pachybrachius). Scudder, 1975. sinae (Rhyparochromus) Stal, 1859.* Two species, Australian region. New Combination (from Pachybrach- INCLUDED SPECIES: ashlockia Scudder, ius). 1975. p. 939.** subgenerica Bergroth, 1918.*** tenua Scudder, 1975. p. 941.*** subinermis Bergroth, 1918.*** DIAGNOSIS: Megapamera is distin- vicina (Nesopamera) Barber, 1958.** guished by its extremely large size (ca. 40 DIAGNOSIS: Pamerana, Megapamera, mm.), being much bigger than any other my- and Afrovertanus are the only myodochine odochine genus. The several fore tibial genera in the Old World with the mesepi- spines found in both sexes is another distin- meron emergent. Afrovertanus, of phallic guishing characteristic. In other myodochine Type IV with a stalklike neck and stridula- genera with spined tibiae, the spines are tory apparatus, is readily separated from found only on male specimens and are usu- Megapamera and Pamerana of phallic Type ally single or few in number. I which both have some head elongation but DESCRIPTION: Body shagreened, ex- lack a neck and are nonstridulatory. Pamer- tremely large and elongate for this tribe of ana is only about one-half the size of its insects (ca. 40 mm.); head elongate with eyes amazingly large sister genus Megapamera protruding; anterior pronotal collar de- and is further distinguished by the charac- marked posteriorly by a linelike groove; lat- teristic median groove in the posterior lip of eral margins of both pronotal lobes rounded. its pygophore. Phallic Type I (fig. 9); conjunctiva and ves- DESCRIPTION: Body elongate; head pro- ica both with areas of greater sclerotization longed behind large, protruding eyes; ante- than that typical of phallic Type I but bearing rior pronotal collar demarked posteriorly by no spines; long, parenthesis-like holding a deep, linelike groove; transverse impres- sclerites typical of Type I present; posterior sion also marked and linelike; lateral margins edge of pygophore broadly rounded (fig. 6); of both pronotal lobes rounded. claval punctation in four or more rows (fig. Phallic Type I (fig. 9); an area of light scler- 17); buccular juncture V-shaped (fig. 81); oc- otization with an irregular edge present on curring at level of antennal insertion; mes- basal portion of vesica, but vesica and con- epimeron emergent (fig. 38); evaporative junctiva both devoid of any actual spines; area extensive (fig. 42) and very rugose; fore long, parenthesis-like holding sclerites typi- femur with spines double-ranked (fig. 15); cal of phallic Type I present; posterior edge fore tibia of both sexes with numerous small of pygophore with a marked deep median stout spines. groove (fig. 8); claval punctation in four or more rows (fig. 17); buccular juncture PAMERANA DISTANT V-shaped (fig. 81); mesepimeron emergent Figure 53 (fig. 38); evaporative area extensive (fig. 42); TYPE SPECIES: Pamerana cuneata Dis- fore femoral spines double-ranked (fig. 15). tant, 1909 =Rhyparochromus nigritulus Walker, 1872. Ten species, Oriental and Palearctic. PACHYBRACHIUS HAHN INCLUDED SPECIES: fulvomaculata Mal- Figure 45 ipatil, 1978. p. 40.*** TYPE SPECIES: Pachybrachius luridus nigritulus (Rhyparochromus) Walker, Hahn, 1826. 1872.** Ten species, Palearctic. 1980 HARRINGTON: MYODOCHINI 91

INCLUDED SPECIES: biguttatus Curtis, a sort of casing close to vesical juncture, but 183-1 .*** no spines present on either conjunctiva or capitatus (Diplonotus) Horvath, 1882.* vesica; long, parenthesis-like holding scler- fasciatus (Plociomerus) Fieber, 1861.*** ites typical of phallic Type I present; poste- festivus (Pamera) Distant, 1883.** rior edge of pygophore subsharp (fig. 5); cla- fracticollis (Pachymerus) Schilling, val punctation in four or more rows (fig. 17); 1829.* buccular juncture V-shaped (fig. 81); mes- fracticollis collaris (Plociomerus) Baer- epimeron enclosed (fig. 37); evaporative area ensprung, 1859. extensive (fig. 42); fore femoral spines dou- fracticollis tridens Roubal, 1959. ble-ranked (fig. 15); male fore tibia with two luridus Hahn, 1826.** tiny spines near distal end. pictus (Pamera) Scott, 1880.*** pusillus (Rhyparochromus) Dallas, 1852, patria ignota.*** STALARIA, NEW GENUS reduviformis Curtis, 1837.*** Figure 47 vaccaroi Mancini, 1954.*** TYPE SPECIES: Pamera ferruginosus Stal, DIAGNOSIS: Pachybrachius, to date, has 1874. been an extremely large (ca. 80 species) Three species, Ethiopian. polyphyletic genus. In this study it has been INCLUDED SPECIES: [All species new recognized that the type species, P. luridus, combinations (from Pachybrachius).] is unlike most of the other myodochines in- ferruginosus (Pamera) Stal, 1874.* cluded in the genus Pachybrachius. Pachy- kisseis (Pachybrachius) Linnavouri, brachius luridus belongs to phallic Type I. 1978. p. 88.* Most of the other species formerly placed in nysias (Pachybrachius) Linnavouri, Pachybrachius belong to phallic Types II 1978. p. 89.* and IV, where they are now recognized as DIAGNOSIS: Stalaria is one of five Old the new genera Bacacephalus, Horridipa- World genera (all of phallic Type I) that have mera, Paraeucosmetus, Pseudopachybrach- a very narrow ringlike collar demarked pos- ius, Pseudoparomius, and Neopamera. teriorly by an extremely deep linelike Pachybrachius, characterized by its minor groove. The collars of Stalaria, Pamerara- autapomorphic variation on phallic Type I, pa, Pamerarma, and Remaudiereana are im- small head slightly elongate behind large punctate and the punctures are so few on the eyes and two tiny spines of the male fore collar of Pachybrachius that they may go tibia, is a small Palearctic complex, including unnoticed. only 10 species. Stalaria, Pamerapa, and Pamerarma have The genus belongs to that group of phallic more elongate bodies than the robust genera Type I with very narrow characteristically Pachybrachius and Remaudiereana. In those ringlike collars (Pachybrachius, Stalaria, three former genera, the total length is great- Pamerapa, Pamerarma, and Remaudi- er than 3.5 times the basal width of the ereana). Pachybrachius still has a few minor pronotum, whereas Pachybrachius and Re- punctures on its collar and the posterior mar- maudiereana have total lengths less than 3.5 gin of the pronotum straight across the base times the width of the posterior pronotal of the scutellum, whereas the other four gen- lobe. era of this group have impunctate collars and Stalaria has numerous long body hairs and sinuate posterior margins of the pronotum. a slender fore femur with single-ranked DESCRIPTION: Body robust, subovoid; spines which serve to distinguish it from Pa- anterior pronotal collar narrow and ringlike, merapa and Pamerarma that lack such long very sparsely punctate, demarked posterior- body hairs and have fore femora with a ly by a deep linelike groove; lateral margins heavy outer median spine in addition to the of both pronotal lobes rounded. single row ranked along the inner edge of the Phallic Type I (fig. 9); an area of light scler- ventral surface. The striking, autapomorph- otization present on conjunctiva and forming ic, multibranched holding sclerites of the 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

male genitalia of Stalaria also characterize long, slender, parenthesis-like holding scler- this new genus. And, unlike the other four ites typical of phallic Type I (these holding genera with ringlike collars, Stalaria has no sclerites are lacking in Pamerarma). spines on the male fore tibia. DESCRIPTION: Body elongate; impunc- DESCRIPTION: Body elongate; impunc- tate anterior pronotal collar very narrow and tate anterior pronotal collar very narrow and ringlike, demarked posteriorly by a deep ringlike, demarked posteriorly by a deep linelike groove; pronotal transverse impres- linelike groove; lateral margins of both pro- sion marked by a similar line; lateral margins notal lobes rounded. of both pronotal lobes rounded. Phallic Type I (fig. 9), but holding sclerites Phallic Type I (fig. 9); a few tiny, hairlike not slender and parenthesis-like, instead spines present on conjunctiva; long paren- heavier and multibranched; posterior edge of thesis-like holding sclerites typical of phallic pygophore with a shallow median groove in Type I present; posterior edge of pygophore lip (fig. 7); claval punctation in four or more with a shallow median groove or depression rows (fig. 17); buccular juncture V-shaped (fig. 7); claval punctation in four or more (fig. 81); mesepimeron enclosed (fig. 37); rows (fig. 17); buccular juncture V-shaped evaporative area extensive (fig. 42); fore (fig. 81); mesepimeron enclosed (fig. 37); femoral spines present only along inner edge evaporative area extensive (fig. 42); fore fe- of ventral surface (fig. 16); male fore tibia mur incrassate with a row of spines along unspined. inner edge of ventral surface and one well- ETYMOLOGY: I am naming this new ge- developed median spine (fig. 14); a single nus for the late C. St'al who described the spine on the curving male fore tibia. type species and whose seminal work with the subfamily Rhyparochrominae laid a groundwork basic to understanding the tribe Myodochini. REMAUDIEREANA HOBERLANDT Figure 52

PAMERAPA MALIPATIL Synonymized with Pachybrachius by Mal- Figure 44 ipatil, 1978, p. 42. Here raised from synon- ymy (see diagnosis for discussion). TYPE SPECIES: Pamera thoracica Dis- TYPE SPECIES: Remaudiereana tibialis tant, 1901. Hoberlandt, 1954. Three species, Australian. Eighteen species, throughout Old World. INCLUDED SPECIES: murrhea (Pamera) INCLUDED SPECIES: [All species new Distant, 1901.** combinations (from Pachybrachius).] pilosa Malipatil, 1978. p. 30.*** africana Hoberlandt, 1954.* thoracica Distant, 1901.** andrewsi (Pamera) Distant, 1901.** DIAGNOSIS: Pamerapa is a member of annulipes (Plociomerus) Baerensprung, the Old World group of phallic Type I with 1859.** narrow, ringlike, impunctate collars. It is boniniensis (Aphanus) Uhler, 1860.* closely and evenly pruinose over the entire castanea (Ptochiomera) Van Duzee, pronotum and the anterior pronotal lobe is 1932.* Transferred to Pachybrachius very enlarged and globose. The male fore tib- by Scudder, 1970. p. 102. ial spine of this genus is also minor in com- flavipes (Plociomerus) Motschulsky, parison to the very large spine on the curving 1863.* fore tibia of Remaudiereana males. horvathi (Diplonotus) Reuter, 1882.* The few tiny conjunctival spines are con- Transferred to Remaudiereana by sidered of independent derivation from the Slater and Wilcox, 1972. p. 957. heavily spined conjunctiva of phallic Type inornatus (Rhyparochromus) Walker, III, as this new genus clearly presents the 1872.* Raised from synonym of Rhy- bulblike, unreduced sperm reservoir and parochromus nigriceps by Scudder, 1980 HARRINGTON: MYODOCHINI 93

1967. p. 269 =palauensis (Pachybrach- punctate anterior pronotal collar very nar- ius) Barber, 1958. Synonymized by row and ringlike, demarked posteriorly by a Malipatil, 1978. p. 50. deep, linelike groove; transverse impression kydippe (Ptochiomera) Kirkaldy, 1905.* also deeply incised; lateral margins of both nigra Scudder, 1963. p. 1236.* pronotal lobes rounded. nigriceps (Rhyparochromus) Dallas, Phallic Type I (fig. 9); posterior edge of 1852.** pygophore with a shallow median groove in noctuabundus (Pamera) Bergroth, lip (fig. 7); claval punctation in four or more 1907.*** rows (fig. 17); buccular juncture V-shaped octonotata (Pamera) Bergroth, 1914.*** (fig. 81); mesepimeron enclosed (fig. 37); puberulus (Orthaea) China, 1930.** evaporative area extensive (fig. 42); fore fe- robustus Malipatil, 1978. p. 55.*** mur with a single row of spines along inner sidnicus (Orthaea) Kirkaldy, 1908.* edge of ventral surface and one well-devel- sobrina (Pamera) Distant, 1901.** oped heavy median spine on outer edge (fig. tibialis (Remaudiereana) Hoberlandt, 14); male fore tibia curving with a single, 1954.** large, well-developed spine in the middle. DIAGNOSIS: Remaudiereana is here res- urrected from synonymy with Pachybrach- PAMERARMA MALIPATIL ius. Pachybrachius is a Palearctic genus dis- Figure 99 tinguished by two small spines apically on the straight male fore tibia, a small head TYPE SPECIES: Orthaea ventralis China, slightly elongate behind the eyes, faint punc- 1930. tures on the anterior pronotal collar, the pos- Two species: Oriental, Australian, Pacific terior margin of the pronotum straight across Oceanic islands. the base of the scutellum and a sclerotized INCLUDED SPECIES: necventralis Mali- conjunctival casing in the male genitalia. Re- patil, 1978. p. 87.*** maudiereana, in contrast, is a widespread ventralis (Orthaea) China, 1930.** genus with many species in the Old World =chinai (Pachybrachius) Usinger, tropics that have a large spine midlength on 1946.** Synonymized by Malipatil, the curving male fore tibia, an impunctate 1978. p. 85. ringlike collar on the anterior pronotal lobe DIAGNOSIS: Pamerarma is one of four and the posterior margin of the pronotum Old World myodochine genera with Type I curving or concave across the base of the male genitalia and a characteristic narrow, scutellum and lack the conjunctival casing of ringlike, completely impunctate collar on the Pachybrachius. anterior pronotal lobe. Within this group it Remaudiereana is one of four Old World, is distinguished from Stalaria, Pamerapa, phallic Type I genera (from ancestor 20) with and Remaudiereana by its small slender narrow, ringlike, completely impunctate col- body form with the head width across the lars. Within this group Remaudiereana is eyes greater than the width of the transverse distinguished by its male genitalia having impression and by the surprising absence of holding sclerites (these are lacking in Pa- holding sclerites in the male genitalia. merarma) and by the lack of conjunctival DESCRIPTION: Body slender, elongate, spines seen in Pamerapa and the lack of parallel-sided; impunctate anterior pronotal elaborate, spatulate, multispined holding collar very narrow and ringlike, demarked sclerites typical of Stalaria. A robust, sub- posteriorly by a deep, linelike groove; trans- ovoid body form and a large spine midlength verse impression also deeply incised; lateral on the curving male fore tibia serve to further margins of both pronotal lobes rounded. identify members of the genus Remaudi- Phallic Type I but with holding sclerites ereana. lacking, conjunctiva and vesica also un- DESCRIPTION: Body robust, subovoid, spined and otherwise unadorned; posterior with many long curving semierect hairs; im- edge of pygophore rounded (fig. 6); claval 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VC)L. 167

punctation with a few punctures in addition edge of pygophore broadly rounded (fig. 6); to three regular rows (fig. 18); buccular junc- claval punctation in four or more rows (fig. ture V-shaped (fig. 81); mesepimeron en- 17); buccular juncture V-shaped (fig. 81); closed (fig. 37); evaporative area extensive mesepimeron enclosed (fig. 37); evaporative (fig. 42); fore femur slender with a single area extensive (fig. 42); fore femur with short row of spines along inner edge of ven- spines double-ranked (fig. 15); male fore tibia tral surface and one moderate spine on outer unspined. edge (fig. 14); male fore tibia relatively straight with a single small spine midlength. PAROMIUS FIEBER Figure 66 MIMOBIUS BERGROTH TYPE SPECIES: Stenocoris gracilis Ram- Figure 75 bur, 1839. TYPE SPECIES: Mimobius capito Ber- Fourteen species, throughout both hemi- groth, 1921. spheres. Two species, Ethiopian. INCLUDED SPECIES: apicatus (Rhyparo- INCLUDED SPECIES: An undescribed chromus) Stal, 1855.* species from Ghana. australis Malipatil, 1978. p. 60.*** capito Bergroth, 1921. No material attenuatus (Rhyparochromus) Dallas, available! 1852.** I have not been able to locate the type dohrnii (Lygaeus (Plociomerus)) Guer- specimen (described from Madagascar) or in, 1857.** any determined material for this genus. For- excelsus Bergroth, 1924.*** tunately, a full dorsal view drawing and a exiguus (Pamera) Distant, 1883.** lateral view of the head were included with gracilis (Stenocoris) Rambur, 1839.* the original description. The appearance of gracilis djoufensis Lindberg, 1938. the insect in those figures is distinctive jejunus (Pamera) Distant, 1883.** enough for me to believe that a series of a limbatus (Pamera) Stal, 1874.* New small ant mimic I examined from Mt. Atewa, Combination (from Pachybrachius). Ghana represents a new species ofMimobius Iongulus (Rhyparochromus) Dallas, which I describe fully elsewhere. The follow- 1852.** ing diagnosis and description are based on pallidus (Plociomerus) Montruzier, those Ghanian specimens. 1865.** DIAGNOSIS: Mimobius is a small ant paraclypeatus Scudder, 1969. p. 174.* mimic (ca. 3.5 mm.). It is distinguished by piratoides (Plociomerus) Costa, 1864.** its general antlike habitus, broad strongly trivialis (Pamera (Paromius)) St'al, declivent head, anterior pronotal lobe shin- 1874.* ing across the calli and distinctive, multi- DIAGNOSIS: Species of Paromius are headed apical conjunctival spines. very elongate with long slender legs and an- DESCRIPTION: Body elongate, highly ant tennae and an abdomen longer than the com- mimetic; head very broad, swollen and de- bined head and pronotal lengths (most my- clivent anteriorly; an anterior collar present odochines have the abdomen shorter than and demarked posteriorly by a linelike the combined lengths of head and prono- groove; pronotum strongly bilobed, anterior tum). A pronotum with the anterior lobe dis- lobe globose; lateral margins of both pro- tinctly lower than the posterior lobe and a notal lobes rounded. characteristic V-shaped collar distinguish Phallic Type II (fig. 10) with the apical con- this genus. The hemelytra are also charac- junctival spines branched and multiheaded; teristic, being uniformly pale and unmarked sawtoothed spiral of minute spines on vesica save for dark punctures and a very minor typical of phallic Type II present; posterior dark area at the corial apex in some species. 1980 HARRINGTON: MYODOCHINI 95

DESCRIPTION: Body elongate and paral- Pseudoparomius, on the other hand, has a lel-sided; pronotum tapering cephalad, an- broad dark transverse band across the hem- terior lobe in a lower plane than posterior elytra and a head width exceeding the width lobe (best observed in lateral view); lateral of the transverse impression. margins of both pronotal lobes rounded; an Pseudoparomius is unique among mem- anterior collar present and demarked poste- bers of phallic Type II in its possession of a riorly by a linelike groove with a character- bifid left apical conjunctival spine. istic median "dip" giving a V-necked ap- DESCRIPTION: Body elongate, head pearance to the collar. broad, little prolonged behind eyes; an an- Phallic Type II (fig. 10); apical conjuncti- terior collar present and demarked poste- val spines slender; posterior edge of pygo- riorly by a linelike groove; anterior pronotal phore broadly rounded (fig. 6); claval punc- lobe not lower than posterior lobe, lateral tation in four or more rows (fig. 17); buccular margins convex; pronotum not dorsoven- juncture V-shaped (fig. 81); mesepimeron en- trally compressed, lateral margins of both closed (fig. 37); evaporative area extensive lobes rounded, not carinate. (fig. 42); fore femur with spines double- Phallic Type II (fig. 10) but with left apical ranked (fig. 15); male fore tibia unspined. conjunctival spine longer than right and bi- fid; posterior edge of pygophore subsharp PSEUDOPAROMIUS, NEW GENUS (fig. 5); claval punctation in four or more Figure 101 rows (fig. 17); buccular juncture V-shaped TYPE SPECIES: Pamera linearis Stal, (fig. 81), close to labial insertion; mesepi- 1874. meron enclosed (fig. 37); evaporative area Two species, Neotropical. extensive (fig. 42); fore femur with spines INCLUDED SPECIES: linearis (Pamera) double-ranked (fig. 15); male fore tibia es- Stal, 1874.* New Combination (from Pachy- sentially straight, unspined. brachius). ETYMOLOGY: From the Greek pseudo, An undescribed species from Brazil and false, for possible confusion of the type Peru. species with members of the genus Parom- DIAGNOSIS: Pseudoparomius is one of ius. only four New World genera with Type II male genitalia. Members of this genus can be distinguished from species of Pseudopachy- PSEUDOPACHYBRACHIUS MALIPATIL brachius and Bacacephalus simply on the Figure 77 basis of size. The latter two genera typically TYPE SPECIES: Rhyparochromus gutta have a total body length less than 4.5 mm., Dallas, 1852. whereas members of Pseudoparomius are Eight species, throughout both hemi- ca. 5.0 mm. or more in length. Pseudopa- spheres. romius linearis exhibits a very slight median INCLUDED SPECIES: [All species but gut- dip in the posterior margin of the anterior tus are new combinations (from Pachybrach- pronotal collar which could lead to this ius).] species being confused with members of the basalis (Rhyparochromus Dallas, genus Paromius in which a V-necked ap- 1852.** pearance of the anterior pronotal collar is capicolus (Pamera) St?al, 1874.* quite characteristic. Yet Paromius has a nar- =dubia (Pamera) Reuter, 1882.* Syn- row head elongate behind the eyes and not onymized by Slater, 1964b. p. 213. as wide across the eyes as the width of the guttus (Rhyparochromus) Dallas, 1852.* pronotal transverse impression, and the de- nesovinctus (Pachybrachius) Ashlock, scribed Western Hemisphere species of Pa- 1972. p. 98.** romius both have entirely pale hemelytra. pacificus (Pamera) Stal, 1874.* =izzardi 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

(Pachybrachius) Hidaka, 1958. Syn- crassiceps (Pamera) Stal, 1874.* New onymized by Scudder, 1970. p. 102. Combination (from Pachybrachius). reductus (Plociomerus) Walker, 1872.* gemmatus (Pamera) Distant, 1918.** undulatus (Plociomerus) Dohrn, 1860.* New Combination (from Pachybrach- vinctus (Pamera) Say, 1831.* ius). DIAGNOSIS: This genus is worldwide in harimaensis (Pamera) Matsumura, distribution and includes several small my- 1913.*** New Combination (from odochines previously of the genus Pachy- Pachybrachius). brachius. Pseudopachybrachius, which in- insignis (Pamera) Distant, 1901.** cludes such species as P. basalis and the leaorum Malipatil, 1978. p. 83.*** ubiquitous species vincta, guttus, and capi- malayus (Rhyparochromus) Stal, 1859.* colus, is best recognized by its small size. New Combination (from Pachybrach- With the exception of the ant mimetic genera ius). Bacacephalus and Mimobius, all other my- mimulus (Pamera) Breddin, 1907.* New odochines of phallic Type II are considerably Combination (from Pachybrachius). larger than the members of Pseudopachy- nervosus (Pamera (Gyndes)) Horvath, brachius. The species ofPseudopachybrach- 1919.* New Combination (from Pach- ius are rather nondescript or undistinguished ybrachius). and are readily separated from Bacacephalus novaeguineae Malipatil, 1978. p. 72.*** and Mimobius, as both of the last two genera pacificus Malipatil, 1978. p. 73.*** are ant mimics with swollen heads and bead- pallicornis (Rhyparochromus) Dallas, like eyes and each has its own unique vari- 1852.** ation from the typical Type II male genitalia papuaguineae Malipatil, 1978, p. 82.*** characteristic of Pseudopachybrachius. perkinsi Malipatil, 1978. p. 75.*** DESCRIPTION: Body small (less than 4.5 sladeni (Pamera) Distant, 1913.** New mm.); lateral margins of both pronotal lobes Combination (from Pachybrachius). rounded; an anterior collar present and de- vitalisi (Pamera) Distant, 1918.** New marked posteriorly by a linelike groove; Combination (from Pachybrachius). hemelytra, in general, largely pale. woodwardi Malipatil, 1978. p. 80.*** Phallic Type II (fig. 10); apical conjuncti- DIAGNOSIS: As discussed by Malipatil val spines slender and dark; posterior edge (1978), the genus Paraeucosmetus is recog- of pygophore broadly rounded (fig. 6); claval nized by a characteristic additional pair of punctation in four or more rows (fig. 17); lobes on the conjunctiva which are not pres- buccular juncture V-shaped (fig. 81); mes- ent in the typical Type II male genitalia. Par- epimeron enclosed (fig. 37); evaporative area aeucosmetus, with 18 species, is the largest extensive (fig. 42); fore femur with spines genus in a complex of 10 genera with Type double-ranked (fig. 15); male fore tibia un- II genitalia. Unfortunately, I have been un- spined. able to find a good external or female syn- apomorphy to distinguish this genus from the PARAEUCOSMETUS MALIPATIL other nine. However, by the following pro- Figure 100 cess of elimination, species of Paraeucos- metus can be identified. TYPE SPECIES: Rhyparochromus palli- Among the myodochine genera with Type cornis Dallas, 1852. II male genitalia, Togo, Eucosmetus, Horri- Eighteen species: Oriental, Ethiopian, dipamera, and Paraparomius have spined Australian, and Pacific Oceanic islands. male fore tibia, whereas Paraeucosmetus INCLUDED SPECIES: albofasciatus does not. Additionally, Paraeucosmetus fe- (Pachybrachius) Barber, 1958.*** New males can be distinguished by wing length Combination (from Pachybrachius). from Togo which is brachypterous, from cincticornis (Rhyparochromus) Walker, Horridipamera and Paraparomius both of 1872.*** which have narrow heads elongate in the 1980 HARRINGTON: MYODOCHINI 97 postocular region, and from Eucosmetus the Two species, Neotropical. species of which have broad heads with INCLUDED SPECIES: An undescribed pronounced supra-antennal jugal ridges like species from Brazil. Paraeucosmetus but have globose, totally globiceps (Pamera) Stal, 1874.* New impunctate anterior pronotal lobes (Paraeu- Combination (from Pachybrachius). cosmetus species have anterior pronotal lobe DIAGNOSIS: This new genus is being rec- punctures at least laterally). Larger size ognized for the Neotropical species Pachy- serves to distinguish Paraeucosmetus species brachius globiceps (Stal). Among the nine from members of Pseudopachybrachius and other genera of phallic Type II, only the Old the two small ant mimetic genera Mimobius World genus Mimobius has a swollen ant and Bacacephalus, all of which generally do mimetic head like that ofBacacephalus. Yet, not exceed 4.5 mm. in total length. Paraeu- the head of Bacacephalus is even more cosmetus species do not exhibit the V- rounded than that of Mimobius and has tiny, necked appearance of the anterior pronotal raised or beadlike eyes. For further identifi- collar or the very elongate body form char- cation, Bacacephalus has basically pale acteristic of Paromius. hemelytra with dark patterning, whereas Of the 10 genera with Type II male geni- Mimobius, in contrast, has dark hemelytra talia, Pseudoparomius is hardest to distin- with pale patterning, and Bacacephalus guish from Paraeucosmetus and one must lacks the spatulate, multispined holding resort to the male genitalia. Pseudoparom- sclerites of Mimobius. ius, a small Neotropical genus of only two In this cladistic analysis, Bacacephalus species, lacks the extra pair of conjunctival has been related to Paraeucosmetus since lobes which characterizes the Old World ge- they both have an extra pair of conjunctival nus Paraeucosmetus but has the apical con- lobes. However, it should be pointed out that junctival spines elongate with the left bifid, these lobes are so distinctive in Bacacephal- whereas these spines are shorter and simple us, looking almost like a lightly sclerotized or not bifid in Paraeucosmetus. second pair of conjunctival spines, that it is DESCRIPTION: Body elongate, parallel- highly possible they represent a convergence sided; head broad with vertex flat and a well- or parallelism rather than a synapomorphy. developed jugal ridge above antennal seg- DESCRIPTION: Body elongate, slender, ment I; an anterior pronotal collar present parallel-sided; head extremely globose, es- and demarked posteriorly by a linelike pecially marked in lateral view; eyes small, groove; lateral margins of both pronotal elevated from head surface, beadlike; an an- lobes rounded; anterior lobe moderately glo- terior pronotal collar present and demarked bose. posteriorly by a linelike groove; transverse Phallic Type II (fig. 10); apical conjuncti- impression deeply incised and anterior pro- val spines symmetrical; an extra pair of lobes notal lobe globose; lateral margins of both midlength on conjunctiva; posterior edge of pronotal lobes rounded, not dorsoventrally pygophore broadly rounded (fig. 6); claval compressed. punctation in four or more rows (fig. 17); Phallic Type II (fig. 10) with an additional buccular juncture V-shaped (fig. 81); mes- pair of spinelike lobes midlength on the con- epimeron enclosed (fig. 37); evaporative area junctiva; apical conjunctival spines slender extensive (fig. 42); fore femur with spines and simple, typical of phallic Type II; pos- double-ranked (fig. 15); male fore tibia terior edge of pygophore rounded (fig. 6); straight, unspined. claval punctation in four or more rows (fig. 17); buccular juncture V-shaped (fig. 81); BACACEPHALUS, NEW GENUS mesepimeron enclosed (fig. 37); evaporative Figure 102 area extensive (fig. 42); fore femur with spines double-ranked (fig. 15); male fore tibia TYPE SPECIES: Pamera globiceps Stal, straight, unspined. 1874. ETYMOLOGY: From the Latin baca, 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 bead, and cephala, head, for the rounded, punctation in four or more rows (fig. 17); beadlike, ant mimetic head of this bug. buccular juncture V-shaped (fig. 81); mes- epimeron enclosed (fig. 37); evaporative area EUCOSMETUS BERGROTH extensive (fig. 42); fore femur with spines Figure 71 double-ranked (fig. 15); male fore tibia es- sentially straight with three or four small TYPE SPECIES: Eucosmetus formosus spines along its distal one-half. Bergroth, 1894. Five species, Oriental. INCLUDED SPECIES: annulicornis Kirit- TOGO BERGROTH shenko, 193 1.** 76 formicarius Breddin, 1907.** Figure formosus Bergroth, 1894.* TYPE SPECIES: Togo victor Bergroth, gessleri Fernando, 1960.*** 1906. incisus (Rhyparochromus) Walker, Three species, endemic, Japan. 1872.** INCLUDED SPECIES: hemipterus (Diplo- DIAGNOSIS: Eucosmetus can be charac- notus) Scott, 1874.** terized by its ant mimetic appearance, broad praetor Bergroth, 1919.*** head with a flattened vertex and prominent victor Bergroth, 1906.*** supra-antennal jugal ridge, extremely globose DIAGNOSIS: Three species are recog- impunctate anterior pronotal lobe, spined nized for Togo, a genus endemic to the is- male fore tibia and characteristic male geni- lands of Japan. All are brachypterous with talia of phallic Type II with asymmetrical an extremely elongate anterior pronotal lobe apical conjunctival spines, the right being (2.5 or more times the length of the posterior short and the left strikingly long and slender. lobe) and curving, spined male fore tibiae. None of the nine other genera of phallic Type Some specimens have a small blunt spine in II have apical conjunctival spines like those the lateral margin of the anterior pronotal of Eucosmetus. Eucosmetus, Horridipa- collar. These collar spines are not so well mera, Paraparomius, and Togo are the only developed as those of the Neotropical genus Type II genera with a spined male fore tibia. Xenydrium and are certainly of independent Paraparomius and Horridipamera have nar- derivation. The bifid right apical conjunctival row heads and Togo is brachypterous, spine of this genus is unique among the gen- whereas Eucosmetus has a broad head and era of phallic Type II. is macropterous. The females of Paraeucos- DESCRIPTrION: Body elongate; head large, metus might be confused with Eucosmetus, broad, slightly prolonged behind eyes; an an- but, on close examination, the entirely im- terior pronotal collar present and demarked punctate, globose anterior pronotal lobe of posteriorly by a linelike groove; pronotum Eucosmetus will serve to identify the genus. strongly bilobed with transverse impression DESCRIPTION: Body elongate, parallel- deeply incised; anterior lobe elongate and sided; head large and broad with flattened convex; lateral margins of both pronotal vertex and marked supra-antennal jugal ridge; lobes rounded. an anterior pronotal collar present and de- Phallic Type II (fig. 10); apical conjuncti- marked posteriorly by a linelike groove; an- val spines asymmetrical, the right larger and terior pronotal lobe globose and impunctate; bifid; posterior edge of pygophore broadly transverse impression deeply incised; lateral rounded (fig. 6); hemelytra typically bra- margins of both pronotal lobes rounded, not chypterous; claval punctation in four or compressed dorsoventrally. more rows (fig. 17); buccular juncture Phallic Type II (fig. 10); apical conjuncti- V-shaped (fig. 81); mesepimeron enclosed val spines asymmetrical, the right short and (fig. 37); evaporative area extensive (fig. 42); the left very long and slender; posterior edge fore femur with spines double-ranked (fig. of pygophore broadly rounded (fig. 6); claval 15); male fore tibia strongly curving with a 1980 HARRINGTON: MYODOCHINI 99 large spine midlength and additional minor with the very broad base to the right apical spines on distal one-half. conjunctival spine (Togo and Eucosmetus also have broad heads). HORRIDIPAMERA MALIPATIL DESCRIPTION: Body elongate, ant mi- Figure 74 metic; head slightly prolonged behind eyes, TYPE SPECIES: Plociomerus nietneri vertex rounded; an anterior pronotal collar Dohrn, 1860. present and demarked posteriorly by a Eleven species, throughout Old World. sharp, linelike groove; pronotum strongly INCLUDED SPECIES: bergrothi (Pamera) bilobed with transverse impression deeply Horvath, 1892.* New Combination (from incised; lateral margins of both pronotal Pachybrachius). lobes rounded. cantrelli Malipatil, 1978. p. 94.*** Phallic Type II (fig. 10); the pair of large ebenaui (Pamera) Reuter, 1887.* New apical conjunctival spines simple, the right Combination (from Pachybrachius). with a very broad base; posterior edge of inconspicuus (Rhyparochromus) Dallas, pygophore broadly rounded (fig. 6); claval 1852.* New Combination (from Pach- punctation in four or more rows (fig. 17); ybrachius). buccular juncture V-shaped (fig. 81); mes- nietneri (Plociomerus) Dohrn, 1860.* epimeron enclosed (fig. 37); evaporative area perlongus (Pachybrachius) Scudder, extensive (fig. 42); fore femur with spines 1971. p. 727.*** New Combination double-ranked (fig. 15); male fore tibia (from Pachybrachius). spined, essentially straight. pullatus (Pamera) Hesse, 1925.* New Combination (from Pachybrachius). PARAPAROMIUS, NEW GENUS robusta Malipatil, 1978. p. 93.*** Figure 73 rusticus (Diplonotus) Scott, 1874.** TYPE SPECIES: Plociomerus leptopoides New Combination (from Pachybrach- Baerensprung, 1859. ius). Two species, Palearctic. spinicrus (Pamera) Reuter, 1882.* New INCLUDED SPECIES: lateralis (Diplono- Combination (from Pachybrachius). tus) Scott, 1874.** New Combination (from subsericeus (Pamera (Entisberus)) Pachybrachius). Breddin, 1907.*** New Combination leptopoides (Plociomerus) Baeren- (from Pachybrachius). sprung, 1859.* New Combination DIAGNOSIS: Horridipamera includes sev- (from Paromius). eral "black and white" ant mimetic species DIAGNOSIS: This new genus is being rec- of phallic Type II that were formerly includ- ognized for two Palearctic species, one here- ed in Pachybrachius. Members of this genus tofore included in the genus Paromius and have a very globose, impunctate anterior one from the genus Pachybrachius. While pronotal lobe with the lines demarking the Paromius lacks the spined male fore tibia of collar and transverse impression both deeply Paraparomius, Paraparomius has the dorsal incised. Eucosmetus, also with a globose, surface of both pronotal lobes in essentially impunctate anterior pronotal lobe, has a the same plane and lacks the V-shaped collar broad head with a flattened vertex. Horridi- and extremely prolonged abdomen of Pa- pamera, in contrast, has a narrow head romius. Paraparomius, like Horridipamera, slightly prolonged behind the eyes and with has a narrow head with a rounded vertex. the vertex very domelike and convex (fig. Both of these genera, as well as Togo and 74). The males of Horridipamera, like those Eucosmetus, have the male fore tibia spined. of Togo, Paraparomius and Eucosmetus Yet, Paraparomius is distinguished by its have the fore tibia spined, but Horridipa- male genitalia with simple, symmetrical api- mera may be distinguished from each of cal conjunctival spines unlike the asymmet- these genera on the basis of its male genitalia rical variations on phallic Type II found in 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 species of Togo, Eucosmetus, and Horridi- metic (field behavior highly ant mimetic); lat- pamera. eral margins of both pronotal lobes rounded, DESCRIPTION: Body elongate, head slen- anterior lobe highly globose and glabrous der, slightly prolonged behind eyes; an an- save collar; a narrow pruinose anterior collar terior pronotal collar present and demarked present and demarked posteriorly by a line- posteriorly by a linelike groove; anterior pro- like groove. notal lobe elongate, not lower than posterior Phallic Type III (fig. 11); posterior edge of pronotal lobe, lateral margins strongly con- pygophore broadly rounded (fig. 6); hemel- vex; lateral margins of both pronotal lobes ytra often brachypterous; claval punctation rounded, not dorsoventrally compressed. in three regular rows (fig. 19); buccular junc- Phallic Type II (fig. 10); conjunctival ture V-shaped (fig. 81); mesepimeron emer- spines short, dark, pointed, and essentially gent (fig. 38); evaporative area extensive (fig. symmetrical; posterior edge of pygophore 42); fore femur with spines only along inner broadly rounded (fig. 6); claval punctation in edge of ventral surface (fig. 16); a lunate, file- four or more rows (fig. 17); buccularjuncture like abdominal stridulitrum present laterally, V-shaped (fig. 81); mesepimeron enclosed extending over sterna II through IV (fig. 35); (fig. 37); evaporative area extensive (fig. 42); plectrum a field of tiny spines or tubercles fore femur with spines double-ranked (fig. on basal one half of hind femur (fig. 36). 15); male fore tibia curving with a large spine ETYMOLOGY: I am pleased to name this midlength and two or three minor spines on new genus for my major dissertation advisor, distal one-half. Dr. James A. Slater, in gratitude for his un- ETYMOLOGY: From the Greek para, be- failing interest, advice, and support of this side or near, for the former placement of the study. type species in the genus Paromius. LIGYROCORIS STAL SLATEROBIUS, NEW GENUS Figure 68 Figure 69 TYPE SPECIES: Cimex sylvestris Lin- TYPE SPECIES: Heraeus insignis Uhler, naeus, 1758. 1872. Eleven species, Nearctic and Neotropical. Two species, Nearctic. INCLUDED SPECIES: balteatus Stal, INCLUDED SPECIES: [Both species are 1874.* new combinations (from Sphaerobius).] caricis Sweet, 1963.** insignis (Heraeus) Uhler, 1872.* delitus Distant, 1882.** quadristriatus (Sphaerobius) Barber, depictus Barber, 1921.** 191 1.** diffusus (Plociomerus) Uhler, 1871 .* DIAGNOSIS: Slaterobius is easily recog- latimarginatus Barber, 1921.** nized as the only genus of phallic Type III litigiosus (Plociomera) Stal, 1862.* with a shining, globose anterior pronotal lobe obscurus Barber, 1921.* contrasting with a pruinose collar and pos- occultus (Pachybrachius) Barber, terior lobe. The only other myodochine with 1953.** New Combination (from this characteristic pattern of pronotal prui- Pachybrachius). nosity is Tenuicoris of phallic Type IV. Ten- slossoni Barber, 1914.** uicoris, however, has an elongate head and sylvestris (Cimex) Linnaeus, 1758. * lacks the stridulatory mechanism exhibited DIAGNOSIS: Ligyrocoris, Slaterobius, and by Slaterobius. Slaterobius is often brachyp- Froeschneria of phallic Type III and Pseu- terous and many specimens with reduced dopamera and Ashlockaria of Type IV have hemelytra also show a peculiar distended abdominal stridulitra extending over the first metapleuron which is very bulbous when three visible sterna and a scattered field of viewed from above. tiny spines or tubercles basally on the hind DESCRIPTION: Body elongate, ant mi- femur as a plectrum. Of these five Nearctic 1980 HARRINGTON: MYODOCHINI 101 genera, only Ligyrocoris and Froeschneria DIAGNOSIS: Only Froeschneria and Li- have the entire dorsal surface of the prono- gyrocoris have the entire dorsal surface of tum pruinose (the pronotum is shining in the pronotum pruinose in combination with Pseudopamera; shagreened in Ashlockaria a stridulatory apparatus like that described and shining on the anterior lobe of Slatero- above. Froeschneria is distinguished from bius). Ligyrocoris by its deeply incised pronotal Ligyrocoris has spines only on the inner transverse impression, annulate fourth an- edge of the ventral surface of the fore femur tennal segment and double-ranked fore fem- (fig. 16) and the transverse impression divid- oral spines. Ligyrocoris has a shallow or ob- ing the two pronotal lobes is shallow, almost solete transverse impression, unicolorous obsolete medially (fig. 68). In Froeschneria fourth antennal segment and spines ranked the fore femoral spines are double-ranked only along the inner edge of the ventral sur- (fig. 15) and the transverse impression is face of the fore femur. deeply incised (fig. 67). DESCRIPTION: Body elongate; lateral DESCRIPTION: Body elongate; lateral margins of both pronotal lobes rounded; margins of both pronotal lobes rounded; an pronotum markedly bilobed with transverse anterior collar present and demarked poste- impression deeply incised; an anterior collar riorly by a linelike groove; transverse pro- present and demarked posteriorly by a line- notal impression shallow. like groove. Phallic Type III (fig. 11); posterior edge of Phallic Type III (fig. 11); conjunctival pygophore broadly rounded (fig. 6); hemel- spines large and numerous; posterior edge of ytra often brachypterous; claval punctation pygophore broadly rounded (fig. 6); claval three regular rows plus a few scattered ad- punctation in four or more rows (fig. 17); ditional punctures (fig. 18); buccularjuncture buccular juncture V-shaped (fig. 81); mes- V-shaped (fig. 81); mesepimeron enclosed epimeron enclosed (fig. 37); evaporative area (fig. 37); evaporative area extensive (fig. 42); extensive (fig. 42); fore femur with spines fore femur with spines only along inner edge double-ranked (fig. 15); antennal segment IV of ventral surface (fig. 16); a lunate, filelike with a pale band basally; a lunate, filelike abdominal stridulitrum present laterally, ex- abdominal stridulitrum extending over sterna tending over sterna II through IV (fig. 35); II through IV laterally (fig. 35); plectrum a plectrum a field of small spines on basal one- field of spines on basal one-half of hind femur half of hind femur (fig. 36). (fig. 36). ETYMOLOGY: I am pleased to name this FROESCHNERIA, NEW GENUS new genus for Dr. Richard C. Froeschner of Figure 67 the United States National Museum of Nat- TYPE SPECIES: Ligyrocoris multispinus ural History. Early in my work with the My- Stal, 1874. odochini, Dr. Froeschner helped to convince Five species, Neotropical (one species me that a generic level revision of the tribe into Nearctic). was the essential first approach. INCLUDED SPECIES: infumatus (Ligyro- coris) Distant, 1882.** New Combination PERIGENES DISTANT (from Ligyrocoris). Figure 72 multispinus Stal, 1874.* New Combina- TYPE SPECIES: Perigenes dispositus Dis- tion (from Ligyrocoris). tant, 1893. oblitus (Ligyrocoris) Distant, 1882.** Two species, Nearctic; one species, Neo- New Combination (from Ligyrocoris). tropical. piligerus (Plociomera) Stal, 1862.* New INCLUDED SPECIES: constrictus (Pa- Combination (from Ligyrocoris). mera) Say, 1831.* vicinalis (Pamera) Distant, 1882.** New dispositus Distant, 1893.** Combination (from Pachybrachius). similis Barber, 1906.** 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

DIAGNOSIS: Perigenes and Zeridoneus genes by stout dark bristles on the tibiae of are the only nonstridulatory myodochines of all legs (Perigenes has no such bristles on the phallic Type III. In both of these genera the fore leg) and a broad, punctate collar unlike sperm reservoir is extremely reduced and the very narrow, ringlike, almost impunctate there are numerous conjunctival spines, in- collar of Perigenes. cluding a group of characteristic very long DESCRIPTION: Body elongate; lateral curving spines and a cap or crownlike assem- margins of both pronotal lobes rounded; blage of spines with a common base. transverse pronotal impression very shallow; The male genitalia of Perigenes show fur- an anterior collar present and demarked pos- ther modification in reduced, truncate clasp- teriorly by a linelike groove. ers (the claspers of Zeridoneus have a more Phallic Type III (fig. 11); sperm reservoir typical curving blade). Perigenes also has a very reduced and elongate; conjunctival very narrow ringlike nearly impunctate col- spines numerous, elongate and in character- lar (fig. 72) unlike the broader punctate collar istic clumps or groups; posterior edge of py- of Zeridoneus (fig. 70). gophore broadly rounded (fig. 6); claval DESCRIPTION: Body elongate robust; lat- punctation in four or more rows (fig. 17); eral margins of both pronotal lobes rounded; buccular juncture V-shaped (fig. 81); mes- a very narrow, sparsely punctate anterior epimeron enclosed (fig. 37); evaporative area collar present and demarked posteriorly by extensive (fig. 42); fore femur with spines a linelike groove. only along inner edge of ventral surface (fig. Phallic Type III (fig. 11); sperm reservoir 16); no stridulatory apparatus present. very reduced and elongate; conjunctival spines numerous, elongate and in character- istic clumps or groups; posterior edge of py- PSEUDOPAMERA DISTANT gophore broadly rounded (fig. 6); claval Figure 94 punctation in four or more rows (fig. 17); buccular juncture V-shaped (fig. 81); mes- TYPE SPECIES: Pseudopamera aurivil- epimeron enclosed (fig. 37); evaporative area liana Distant, 1882. extensive (fig. 42); fore femur with a few Eight species, Nearctic and Neotropical. spines only along inner edge of ventral sur- INCLUDED SPECIES: ater (Prytanes) Dis- face (fig. 16); no stridulatory apparatus pres- tant, 1893.** New Combination (from Pry- ent. tanes). aurivilliana Distant, 1882.* ZERIDONEUS BARBER coloradensis (Ligyrocoris (Neoligyro- Figure 70 coris)) Barber, 1921. ** insititius (Erlacda) Distant, 1893.** TYPE SPECIES: Perigenes costalis Van New Combination (from Ligyrocoris). Duzee, 1909. nitidicollis (Pamera) Stal, 1874.* Three species, Nearctic. nitidula (Pamera) Uhler, 1893.** INCLUDED SPECIES: costalis Van Duzee, rubricata (Ligyrocoris (Neoligyrocoris)) 1909. * Barber, 1921.** knulli Barber, 1948.* setosa (Pamera) Stal, 1874.** New petersoni Reichart, 1966. p. 347.*** Combination (from Ligyrocoris). DIAGNOSIS: As indicated in the preced- DIAGNOSIS: Pseudopamera is one of ing diagnosis for Perigenes, Zeridoneus and eight myodochine genera with an abdominal Perigenes lack a stridulatory apparatus and stridulitrum. This type of stridulatory mech- have phallic Type III with greater sperm res- anism is found in Stridulocoris with Type I ervoir reduction and conjunctival spine elab- male genitalia, Slaterobius, Ligyrocoris, and oration than the three stridulatory members Froeschneria of Type III and Erlacda, Af- of this phallic type. rovertanus, Ashlockaria, and Pseudopamera Zeridoneus is distinguished from Peri- which are "Type IV bugs." 1980 HARRINGTON: MYODOCHINI 103

Pseudopamera may readily be distin- INCLUDED SPECIES: arhaphaeoides Sig- guished from Erlacda and Afrovertanus by noret, 1863.* its plectrum. In the latter two genera the signoreti Porter, 1929.* plectrum consists of two or three chisel-like DIAGNOSIS: This tiny ant mimetic genus projections at the base of the hind femur, is distinguished by its relatively short ab- whereas in Pseudopamera it is a fairly ex- dominal stridulitrum and chisel-like plec- tensive field of small spines on the hind fe- trum. Only the Old World genus Afrovertan- mur. Ashlockaria (also of Type IV) has a us, with a longer neck and the mesepimeron plectrum similar to that of Pseudopamera, emergent, has a plectrum like that of Erlac- but Ashlockaria has a shagreened body sur- da. Some male Erlacda specimens addition- face texture and multiple male fore tibial ally show a distinctive pair of small tubercles spines while Pseudopamera is shining with or horns on the head behind the ocelli. a single spine on the male fore tibia. DESCRIPTION: Body elongate, shining, Pseudopamera may also be distinguished very ant mimetic; head prolonged behind from Stridulocoris, Slaterobius, Ligyrocoris, eyes in a short but distinct neck; lateral mar- and Froeschneria by its shining dorsum. The gins of both pronotal lobes rounded; an an- dorsal surface of the pronotum is pruinose in terior pronotal collar present and demarked Ligyrocoris, Froeschneria, and Stridulocoris posteriorly by a linelike groove. (although the pruinosity is made less appar- Phallic Type IV (fig. 12); posterior edge of ent by heavy punctation in Stridulocoris). In pygophore broadly rounded (fig. 6); hemel- Slaterobius the anterior pronotal lobe is shin- ytra often brachypterous; claval punctation ing but the collar and posterior pronotal lobe in three regular rows (fig. 19); buccular junc- are characteristically pruinose. ture V-shaped (fig. 81) and extended poste- DESCRIPTION: Body elongate, parallel- riorly in a slight midventral carina; mesepi- sided, shining; head slightly elongate behind meron enclosed (fig. 37); evaporative area eyes; pronotal lobes rounded; an anterior extensive (fig. 42); fore femur multispined as collar present and demarked posteriorly by in figure 13; male fore tibia with a small spine a linelike groove. on distal one-half; a short, broadly V-shaped, Phallic Type IV (fig. 12); posterior edge of filelike abdominal stridulitrum extending pygophore broadly rounded (fig. 6); claval over the lateral surface of sterna II and III punctation in four or more rows (fig. 17); (fig. 28); plectrum three chisel-like projec- ventral surface of head not grooved; buccu- tions at base of hind femur (fig. 29). lar juncture V-shaped (fig. 81), mesepimeron here enclosed (fig. 37); evaporative area re- AFROVERTANUS SCUDDER duced (fig. 41); fore femur with spines dou- Figure 88 ble-ranked (fig. 15); male fore tibia with a spine on distal one-half; a lunate, filelike ab- TYPE SPECIES: Afrovertanus elongatus dominal stridulitrum present, extending lat- Scudder, 1962.* erally over sterna II through IV (fig. 35); Monotypic, Ethiopian. plectrum a field of small spines on base of DIAGNOSIS: Afrovertanus is the only my- hind femur (fig. 36). odochine with phallic Type IV occurring in the Old World. Only Afrovertanus and Ae- gyptocoris among the Eastern Hemisphere ERLACDA SIGNORET genera have the head prolonged posteriorly Figure 85 in a stalklike neck and the dorsal body sur- face highly shining. Afrovertanus may be dis- TYPE SPECIES: Erlacda arhaphaeoides tinguished from Aegyptocoris by its emer- Signoret, 1863. * gent mesepimeron, abdominal stridulitrum, =Sphaerobius gracilis Uhler, 1893.** and by the lack of an apically upturned scu- NEW SYNONYMY. tellum which is characteristic of Aegypto- Two species, Neotropical. coris. 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167

DESCRIPTION. Body elongate, shining; XENYDRIUM POPPIUS AND BERGROTH head prolonged behind eyes in a stalklike Figure 92 neck; lateral margins of both pronotal lobes TYPE SPECIES: Xenydrium formiciforme rounded; an anterior pronotal collar present Bergroth, 1921.* and demarked posteriorly by a linelike Monotypic, Neotropical. groove. DIAGNOSIS: Xenydrium is readily recog- Phallic Type IV (fig. 12); posterior edge of nized by its strikingly ant mimetic habitus pygophore sharp (fig. 4); claval punctation in with the antenniferous tubercles broadly ex- four or more rows (fig. 17); buccularjuncture panded in shelflike plates that are very man- V-shaped (fig. 81); mesepimeron emergent dibular in appearance. The extreme anterior (fig. 38); evaporative area reduced (fig. 41); narrowing of the abdomen also enhances this fore femur with spines single-ranked, present mimicry. The small but distinctive spines on only along inner edge of ventral surface (fig. the lateral margins of the collar are charac- 16); a distinctive lunate, filelike abdominal teristic for the genus. stridulitrum present laterally on sterna II DESCRIPTION: Body elongate, shining, through IV (fig. 33); plectrum two chisel-like and strikingly ant mimetic; head especially projections at base of hind femur (fig. 34). antlike with a very short neck region behind eyes; lateral margins of both pronotal lobes rounded; an anterior collar present and de- PSEUDOCNEMODUS BARBER marked posteriorly by a linelike groove; a Figure 84 small spine present on the lateral margin of TYPE SPECIES: Pseudocnemodus bruneri the collar. Barber, 1911.** Phallic Type IV (fig. 12) posterior edge of =Pterotmetus canadensis Provancher, pygophore subsharp (fig. 5); hemelytra often 1886. brachypterous; claval punctation in four or Monotypic, Nearctic. more rows (fig. 17); buccular juncture DIAGNOSIS: Pseudocnemodus can be V-shaped (fig. 81); mesepimeron enclosed recognized readily by its unique stridulatory (fig. 37); evaporative area extensive (fig. 42); mechanism. The stridulitrum consists of arc- fore femur with spines double-ranked (fig. like, cross-striated areas on the propleuron 15); male fore tibia with a well-developed and lateral surface of the head (fig. 39), spine. whereas the plectrum is a row of tubercles on the proximal one third of the fore femur ASHLOCKARIA, NEW GENUS (fig. 40). Figure 90 DESCRIPTION: Body elongate, sha- TYPE SPECIES: Cnemodus sobrius Uhler, greened; head elongate behind eyes but lack- 1894. ing a distinct neck region; lateral margins of Two species, Nearctic. both pronotal lobes rounded; an anterior col- INCLUDED SPECIES: An undescribed lar present and demarked posteriorly by a species from California. linelike groove. sobrius (Cnemodus) Uhler, 1894.* New Phallic Type IV (fig. 12); posterior edge of Combination (from Pseudopamera). pygophore subsharp (fig. 5); hemelytra often DIAGNOSIS: Ashlockaria is one of three brachypterous; claval punctation in four or genera of phallic Type IV with an exten- more rows (fig. 17); buccular juncture sively shagreened body surface. With Zero- V-shaped (fig. 81); mesepimeron enclosed pamera it also shares a multispined male fore (fig. 37); evaporative area reduced (fig. 41); tibia and peculiar elongate curving body fore femur with spines double-ranked (fig. hairs on the legs as well as the head and 15); male fore tibia with a well-developed pronotum. Cnemodus, the third shagreened spine. myodochine with a Type IV phallus, lacks 1980 HARRINGTON: MYODOCHINI 105

such hairs and multiple male fore tibial no light areas patterning the body or hemel- spines and also lacks ocelli. Both Ashlock- ytra. Cnemodus also has a largely dark body, aria and Zeropamera are large and rather but in Cnemodus the legs and antennae are coleopteroid in appearance (presumably ant pale, and Cnemodus is also readily recog- mimetic in behavior), but only Ashlockaria nized by its lack of ocelli. has an abdominal stridulitrum. Thus, it is DESCRIPTION: Body elongate, sha- distinct as the only stridulatory myodochine greened, with numerous long curving hairs; with a shagreened body surface. On the basis lateral margins of both pronotal lobes round- of its independently derived stridulatory ed; an anterior collar present and demarked mechanism, the former Pseudopamera sob- posteriorly by a linelike groove. rius has been given generic status. Phallic Type IV (fig. 12); posterior edge of DESCRIPTION: Body elongate, sha- pygophore broadly rounded (fig. 6); claval greened, with numerous long curving hairs; punctation in four or more rows (fig. 17); lateral margins of both pronotal lobes round- buccular juncture V-shaped (fig. 81); mes- ed; an anterior collar present and demarked epimeron enclosed (fig. 37); evaporative area posteriorly by a linelike groove. extensive (fig. 42); fore femur with spines Phallic Type IV (fig. 12); posterior edge of double-ranked (fig. 15); male fore tibia mul- pygophore broadly rounded (fig. 6); hemel- tispined. ytra sometimes brachypterous; claval punc- tation in four or more rows (fig. 17); buccular CNEMODUS HERRICH-SCHAEFFER juncture V-shaped (fig. 81); mesepimeron en- Figure 98 closed (fig. 37); evaporative area extensive (fig. 42); fore femur with spines double- TYPE SPECIES: Cnemodus brevipennis ranked (fig. 15); male fore tibia multispined; Herrich-Schaeffer, 1850. a filelike abdominal stridulitrum present lat- =Astemma mavortia Say, 1831. erally on sterna II through IV, though some- Three species, Nearctic; two species (?), times difficult to discern; plectrum a field of Neotropical. spines on base of hind femur. INCLUDED SPECIES: albimaculus Berg, ETYMOLOGY: I am pleased to name Ash- 1879.*** lockaria for Dr. Peter Ashlock of the Uni- brevipennis Herrich-Schaeffer, 1850.* versity of Kansas whose preliminary work hirtipes Blatchley, 1924.* and continuing interest in the Myodochini, inflatus Van Duzee, 1915.* as well as the loan of many specimens, have multifarius Berg, 1894.*** enhanced this study. DIAGNOSIS: Many specimens of Cnemo- dus are coleopteroid. In the field these in- ZEROPAMERA BARBER sects are quite ant mimetic by virtue of their Figure 95 behavior. The genus has a shagreened body surface like that of its near relatives, Ash- TYPE SPECIES: Zeropamera nigra Bar- lockaria and Zeropamera, but is easily dis- ber, 1948.** tinguished from those two genera and from Monotypic, Nearctic. all other myodochine genera by its lack of DIAGNOSIS: Zeropamera has a sha- ocelli. greened body surface like that of Ashlock- DESCRIPTION: Body elongate, sha- aria and Cnemodus and additionally has the greened; lateral margins of both pronotal long body hairs and multispined male fore lobes rounded; an anterior collar present and tibiae ofAshlockaria. It lacks the abdominal demarked posteriorly by a linelike groove. stridulitrum of Ashlockaria and is best dis- Phallic Type IV (fig. 12); posterior edge of tinguished by its large size and dark color- pygophore broadly rounded (fig. 6); hemel- ation. Zeropamera is uniformly an extremely ytra sometimes brachypterous or subma- dark chestnut brown verging on black with cropterous; claval punctation in four or more 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 rows (fig. 17); buccular juncture V-shaped oval eyes and a midventral carina prolonged (fig. 81); mesepimeron enclosed (fig. 37); from the buccular juncture as does Disting- evaporative area extensive (fig. 42); fore fe- physes. Yet Pephysena is distinguished by a mur with spines double-ranked (fig. 15); male longer neck and an emergent mesepimeron fore tibia with a single spine. not seen in Distingphyses. DESCRIPTION: Body elongate, ant mi- DISTINGPHYSES SCUDDER metic; head elongate with a distinct stalklike Figure 89 neck; eyes elongate, oval; lateral margins of TYPE SPECIES: Pephysena insignis Dis- both pronotal lobes rounded; an anterior col- tant, 1882.** lar present and demarked posteriorly by a Monotypic, Neotropical. linelike groove. DIAGNOSIS: This monotypic genus has a Phallic Type IV (fig. 12); posterior edge of heavily punctate ant mimetic head (fig. 89) pygophore broadly rounded (fig. 6); claval and oval eyes (like Pephysena). However, punctation in four or more rows (fig. 17); the neck of Distingphyses is much less ap- buccular juncture V-shaped (fig. 81), con- parent than that of Pephysena and Disting- tinuing posteriorly as a strong midventral ca- physes is further distinguished by a shining rina; mesepimeron emergent (fig. 38); evap- anterior pronotal lobe, including the collar. orative area extensive (fig. 42); fore femur DESCRIPTION: Body elongate, ant mi- with spines double-ranked (fig. 15); male fore metic; head very antlike, elongate behind tibia unspined. eyes with a short barely discernible neck; eyes elongate, oval; lateral margins of both TENUICORIS SLATER AND HARRINGTON pronotal lobes rounded; transverse impres- Figure 96 sion deeply incised; an anterior collar pres- ent and demarked posteriorly by a linelike TYPE SPECIES: Tenuicoris myrmeforme groove. Slater and Harrington, 1974. p. 174.** Phallic Type IV (fig. 12); posterior edge of Monotypic, Neotropical. pygophore broadly rounded (fig. 6); claval DIAGNOSIS: Tenuicoris is readily distin- punctation in four or more rows (fig. 17); guished by its globose, highly shining, im- buccular juncture V-shaped (fig. 81), con- punctate anterior pronotal lobe which con- tinuing posteriorly as a strong midventral ca- trasts with a punctate, pruinose anterior rina; mesepimeron enclosed (fig. 37); evap- collar and posterior lobe. The only other orative area extensive (fig. 42); fore femur myodochine genus with such a pruinosity with spines double-ranked (fig. 15); male fore pattern is Slaterobius which is stridulatory tibia unspined. and of a different phallic type. The head of Tenuicoris is also quite dis- PEPHYSENA DISTANT tinctive with a flattened vertex and the juga Figure 86 forming expanded ridges above the first an- tennal segments (fig. 96). TYPE SPECIES: Pephysena levis Distant, DESCRIPTION: Body very elongate, slen- 1882. der, and ant mimetic; legs and antennae also Five species, Neotropical. extremely long and slender; head prolonged INCLUDED SPECIES: fuscosa Barber, behind eyes, forming a neck region; vertex 1954.* of head very flat, depressed between eyes; levis Distant, 1882.** eyes elongate, oval; lateral margins of both picta Barber, 1954.** pronotal lobes rounded; a narrow anterior An undescribed species from Brazil. collar present and demarked posteriorly by An undescribed species from Surinam. a linelike groove. DIAGNOSIS: Pephysena has a heavily Phallic Type IV (fig. 12); posterior edge of punctate, ant mimetic head with elongate pygophore broadly rounded (fig. 6); claval 1980 HARRINGTON: MYODOCHINI 107

punctation in four or more rows (fig. 17); no satisfactory synapomorphic character buccular juncture V-shaped (fig. 81), con- state to distinguish the group, which may tinuing posteriorly as a strong midventral ca- prove on closer species level examination rina; mesepimeron emergent (fig. 38); evap- not to be holophyletic. orative area extensive (fig. 42); fore femur Neopamera belongs to the Neotropical very slender with only a few spines present element of phallic Type IV in which the on both inner and outer edges of ventral sur- members have a pruinose body surface and face. unspined male fore tibia (from common ancestor 44 in the cladogram). Within this there are NEOPAMERA, NEW GENUS group, various degrees of head Figure 83 elongation, but all the species ofNeopamera show little if any elongation of the head be- TYPE SPECIES: Pamera bilobata Say, hind the eyes and certainly no neck region 1831. such as that exhibited by most other mem- Twenty species: predominantly Neotropi- bers of the group. This Neotropical element, cal, five Nearctic. in addition to exhibiting head elongation, is INCLUDED SPECIES: [All 20 species are characterized by long legs and antennae. new combinations (from Pachybrachius).] These insects are rapidly running forms. In albocinctus (Pachybrachius) Barber, Neopamera, and, in fact, in all but Disting- 1953.** physes, the mesepimeron is emergent. The bilobata (Pamera) Say, 1831.* emergent mesepimeron may be used to dis- brachialis (Rhyparochromus (Plociom- tinguish all Western Hemisphere species cur- erus) Stal, 1858.** rently included in Pachybrachius that are costalis (Pamera) Stal, 1874.* here being recognized as constituting the crassicornis (Pamera) Stoal, 1874.*** new genus Neopamera (note exception of honduranus (Pamera) Bergroth, 1914.* the following monotypic genus Orthaea insularis (Orthaea) Barber, 1925.** which is being resurrected to generic status). intermedius (Orthaea) Barber, 1924. * Among New World myodochines with a mumfordi (Ptochiomera) Van Duzee, pruinose body surface, unspined fore tibia, 1935.*** Transferred to Pachybrach- and phallic Type IV, Neopamera can be dis- ius by Scudder, 1970. p. 102. tinguished from the complex from ancestor neotropicalis (Orthaea) Kirkaldy, 1909.* 47 (Orthaea, Catenes, Heraeus, Myodocha) paganus (Pamera) White, 1879.*** by virtue of its head with a flattened vertex platanus (Pamera) Bergroth, 1894.*** and not prolonged behind the eyes. The near procerulus (Pamera) Berg, 1892.*** relatives Distingphyses, Pephysena, and Ten- recinctus (Pamera) Breddin, 1901.** uicoris have flattened heads like Neopa- serripes (Rhyparochromus) Walker, mera, but all three of the former genera have 1872.* Transferred to Pachybrachius some degree of head elongation and elongate by Scudder, 1970. p. 102. oval eyes, while Neopamera has round eyes sororculus (Pamera) Berg, 1892.*** and the head not prolonged. tineodes (Pachymerus) Burmeister, DESCRIPTION: Body elongate; head not 1835.*** elongate, vertex flat; lateral margins of both tuberculatus (Pamera) Osborn, 1904.*** pronotal lobes rounded; an anterior collar vicarius (Pachybrachius) Barber, 1954.** present and demarked posteriorly by a line- vividus (Pamera) Distant, 1882.** like groove. DIAGNOSIS: This large new genus is Phallic Type IV (fig. 12); posterior edge of being erected to include all the species of a pygophore broadly rounded (fig. 6); claval New World, essentially Neotropical com- punctation in four or more rows (fig. 17); plex, heretofore included in the genus buccular juncture V-shaped (fig. 81); mes- Pachybrachius. Unfortunately, I have found epimeron emergent (fig. 38); evaporative 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 area extensive (fig. 42); fore femur with punctation in four or more rows (fig. 17); spines double-ranked (fig. 15); male fore tibia buccular juncture V-shaped (fig. 81); mes- unspined. epimeron emergent (fig. 38); evaporative ETYMOLOGY: From the Greek neo-, area extensive (fig. 42), extremely rugose; new, Pamera, for an old generic (subgeneric) fore femur slender with few spines but those name by which many of the species of Neo- present ranked along both inner and outer pamera were formerly known. edges of ventral surface; male fore tibia un- spined. ORTHAEA DALLAS Figure 91 CATENES DISTANT Here raised from synonymy with Pachy- Figure 93 brachius. TYPE SPECIES: Catenes porrectus Dis- TYPE SPECIES: Orthaea consuta Dallas, tant, 1893.** 1852. Monotypic, Neotropical. Two species, Neotropical. DIAGNOSIS: This monotypic genus has a INCLUDED SPECIES: [Both species are distinctive elongate flattened head (fig. 93). new combinations (from Pachybrachius).] The reduction of fore leg armature accom- consuta Dallas, 1852.** panied by dark spots of seemingly heavy procinctus (Pamera) Breddin, 1901.* sclerotization which give a "freckled" ap- DIAGNOSIS: Orthaea consuta is resur- pearance to otherwise pale slender fore fem- rected to generic status and Pachybrachius ora is also characteristic of C. porrectus. procinctus (Breddin) is transferred to Or- DESCRIPTION: Body elongate; head pro- thaea. The habitus of these two species is longed behind eyes but with lateral margins unlike that of the other New World "Pachy- nearly parallel, not constricted to a stalklike brachius" species with which they have neck; lateral margins of both pronotal lobes been associated. The small slightly elongate rounded; an anterior collar present and bare- head placed in a noticeably lower plane than ly demarked posteriorly by a linelike groove. the posterior pronotal lobe is very charac- Phallic Type IV (fig. 12); posterior edge of teristic. The fore leg armature of Orthaea is pygophore broadly rounded (fig. 6); claval very reduced and some specimens examined punctation in four or more rows (fig. 17); even lack a fore coxal spine (Neopamera buccular juncture V-shaped (fig. 81); mes- species have generally one and sometimes epimeron emergent (fig. 38); evaporative two well-developed fore coxal spines and area extensive (fig. 42); fore femur with numerous double-ranked fore femoral spines only along inner edge of ventral sur- spines). face (fig. 16). Orthaea is also readily distinguished by a uniformly blackish brown body with distinc- tive bright orange markings medially and on HERAEUS STAL the lateral margins of the posterior pronotal Figure 87 lobe and subapically on the corium. No other TYPE SPECIES: Lygaeus (Plociomerus) described myodochines are similarly pig- triguttatus Guerin, 1857. mented. Twelve species, Neotropical and Nearctic. DESCRIPTION: Body elongate; head with INCLUDED SPECIES: cincticornis St'al, a rounded vertex, slightly prolonged behind 1874.* eyes but not constricted in a neck; lateral cinnamomeus Barber, 1948.** margins of both pronotal lobes rounded; an coquilletti Barber, 1914.* anterior collar present and faintly demarked elegans (Nabis) Walker, 1873.*** Raised by a linelike groove. from synonymy with Heraeus gutta- Phallic Type IV (fig. 12); posterior edge of tus Dallas, 1852 by Scudder, 1967. p. pygophore broadly rounded (fig. 6); claval 264. 1980 HARRINGTON: MYODOCHINI 109

FIG. 103. Cladogram of Genera of Myodochini: Type numbers at bottom indicate male genitalic type for all the genera in the area above that type number. Small numbers in grid at top of cladogram indicate the number of species known for the genus in a particular zoogeographic region.

eximius Distant, 1882.** the head is very slender and rounded, being guttatus (Orthaea) Dallas, 1852.** visually not of much greater diameter than illitus Distant, 1882.** the very elongate neck (fig. 97). In Heraeus pacificus Barber, 1925.* (fig. 87) the head is rounded and far more plebejus Stal, 1874.* bulbous on the end of a stalklike neck which pulchellus Barber, 1954.** is relatively shorter than the neck of Myodo- triguttatus [Lygaeus (Plociomerus)] cha. The anterior pronotal collar of Heraeus Guerin, 1857.* is distinctive, being very narrow dorsally but variegatus Kirby, 1890.*** broad and extending forward beneath the DIAGNOSIS: Heraeus belongs to the Neo- head ventrally. tropical pruinose phallic Type IV group with DESCRIPTION: Body elongate; head pro- the male fore tibia unspined (from common longed behind eyes and constricted to form a ancestor 47 in cladogram). Within that stalklike neck; lateral margins of both pro- group, Heraeus and Myodocha have the notal lobes rounded; an anterior collar pres- longest most stalklike necks. In Myodocha ent and demarked posteriorly by a linelike 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 167 groove, ventrally extending forward beneath DIAGNOSIS: Myodocha is distinguished the head. by the greatest degree of head elongation in Phallic Type IV (fig. 12); posterior edge of the tribe Myodochini. The head of Myodo- pygophore broadly rounded (fig. 6); claval cha is characteristically highly shining, in punctation in four or more rows (fig. 17); contrast with a heavily pruinose pronotum, buccular juncture V-shaped (fig. 81); mes- and very slender in the ocular and preocular epimeron emergent (fig. 38); evaporative areas as well as in the stalklike neck region area extensive (fig. 42); fore femur with (fig. 97). spines double-ranked (fig. 15). DESCRIPTION: Body legs and antennae very elongate; head also long and slender, MYODOCHA LATREILLE strikingly prolonged behind eyes as a stalk- Figure 97 like neck, anterior portion of head visually of little greater diameter than neck region; TYPE SPECIES: Myodocha serripes Oli- lateral margins of both pronotal lobes round- vier, 1811, by action of International Com- ed; a narrow anterior collar present and mission Zoological Nomenclature, Opinion faintly demarked posteriorly by a linelike 669. groove. Eight species, Neotropical and Nearctic. IV of INCLUDED SPECIES: annulicornis Blatch- Phallic Type (fig. 12); posterior edge ley, 1926.* pygophore broadly rounded (fig. 6); claval fulvosa Barber, 1954.** punctation in four or more rows (fig. 17); giraffa Stal, 1862.* buccular juncture V-shaped (fig. 81); mes- inermiba Distant, 1882.** epimeron emergent (fig. 38); evaporative intermedia Distant, 1882.** area extensive (fig. 42); fore femur with longicollis Stal, 1874.* spines double-ranked (fig. 15). serripes Olivier, 1811.* unispinosa Stal, 1874.*

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1932. New Hemiptera-Heteroptera from the 1873. Catalogue of Hemiptera Heteroptera in Marquesas. Bull. Bishop Mus., vol. 98, the British Museum, Supplement. Lon- pp. 177-191. don, E. W. Janson, 63 pp. 1935. Note on Ptochiomera caeca Van Du- White, F. B. zee. Pan-Pacif. Ent., vol. 11, p. 174. 1879. Descriptions of new Hemiptera (I). Walker, F. Jour. Linn. Soc. (Zool.), vol. 14, pp. 1872. Catalogue of the specimens of Hemip- 482-489. tera Heteroptera in the collection of the British Museum. London, 8 parts. .,,,_,, 11 -... ,,.,7x"l-Ki ".w jk.- -%,Z-'.-, -, ,-;,',.t. ', 1- .. .1_11 1-11, "..7* !14 , 7 -` .lw.__..,.1I--.. , ,"',_ .1.1'.'. SF, ,i- I:," . .-.1..l ,4 F,;II",'',..111, IN 1,- ..;) ,I!,:,__J...!-zi,.,-,","', M,P,! .1, ,.-.-- 4, ., :. .l-.` .., ._ ., .4 _..:, "I'M,XI, ,7 .,,1,-",-. ,. :'. _11,4 ;. R!,._b ,.,,.,"I Ir ..'L,.. ,.. ..,-I-,i,...,.,I`.-1.-1).,__".I5.19, .l"I,,,,..11.".-,..-.. ll.. 4T-wrz-7, .,...I,...;,,-I-,.,,.-"..-t,r..,,."..""...11,,; '",,T..-,-- .'. r. ..._ ". ,. 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