Version 2.2 of the Plant Bugs of the World (Insecta: Heteroptera: Miridae) on-line database is based on the same MySQL schema as version 2.0, but offers a more flexible approach to querying the data via the inclusion of geographic searching, as well as offering some other additional enhancements and corrections. Queries are organized around browsing the taxonomic hierarchy or the use of search boxes.
Browse the Taxonomic Hierarchy
Search Boxes allow the use of two strategies
Scientific name searches:
Additional Search Features
Hierarchic display of taxonomic information:
All taxonomic information is displayed in a hierarchic format. To proceed to the next level in the hierarchy choose the green arrow. Scientific names and author names serve as links to additional information about those names. Names of genera and species are listed alphabetically under the next higher categorical level in the database. Junior synonyms are listed under the senior name. Thus, if you are searching for a species (say marmoratus) in a large genus such as Phytocoris, but do not know if the name is currently valid, you may wish to query that species name and select from the list that is returned; to discover junior synonyms, query using “sci. name pattern”. When viewing references for a valid name, entries for the senior synonym and its junior synonym(s) are returned. When viewing references for a junior synonym, only references for the junior name are returned.
Names as links:
Green arrows as links:
Digital Library access:
Digital library entries are available for publications dealing with the Orthotylinae and Phylinae where permission to use those papers on the Internet has been secured. Digital library information can be accessed in three ways:
NOTE: The literature search for this version of the catalog in reasonably complete through 2010, adding approximately eight years to the literature covered in version 2.1. A few references currently seen in the Zoological Record, as of August 2011, are still missing from the catalog; taxa described or mentioned in them will be added when copies of the papers become available.
Some aspects of data in the catalog are still in need of work. Among these are the indications of type species. Most genera have type species indicated and most of these are correct. Nonetheless, some type species indications are currently missing, and a few are incorrect. This situation is in the process of being remedied.
Minor enhancements to the interface are also underway.
This database is derived from Plant Bugs of the World (Schuh, 1995), a work that was heavily based on J. C. M. Carvalho's monumental Catalogue of the Miridae of the World, published between 1957 and 1960. Carvalho had surveyed the literature from 1758 through 1955; Schuh (1995) added data from 1956--1993. The present work adds additional literature and offers the possibility for continued updating; it also adds enhancements to the underlying database and searching functions. As with the work of Schuh (1995), this database does not include entries for all references to all species published before 1956, but for the most part includes only references for that period in which names were made available. In addition it points to the volume and page in Carvalho's work where additional pre-1956 literature references might be found. Those catalog pages can now be viewed through the digital library for the subfamilies Orthotylinae and Phylinae. Corrections to the 1995-version of the catalog as identified and published by Kerzhner and Schuh (2001) are included in the database, as are most new names and other nomenclatural changes published between 1993 and 2011. Most entries are based on direct examination of the primary literature.
Schuh (1995) pointed out that in the 40 years intervening between the publication of the Carvalho Catalog and his own successor to it, that Carvalho's work had become badly outdated for several reasons. Although the period between the appearance of this database online and the publication of Schuh's work is much shorter than that between the works of Carvalho and Schuh, the numbers of additions and changes to the taxonomy of the Miridae are nonetheless remarkable. The following tables and graph provide data on historical growth in our knowledge of the Miridae.
Table 1. Numbers of Available Names
|Described before 1956||1,145|
|Described between 1955 and 1994||678|
|Described between 1993 and 2002||120|
|Described between 2001 and 2011||122|
|Described before 1956||6,637|
|Described between 1955 and 1994||5,465|
|Described after 1993 and 2002||945|
|Described between 2001 and 2011||910|
Table 2. Numbers of Papers Published
|Cataloged papers published before 1956||1307|
|Papers cataloged published between 1956-1993||1927|
|Papers cataloged published between 1993 and 2002||354|
|Papers cataloged published between 2001 and 2011||263|
Table 3. Numbers of species described, by decade.
Table 4. Numbers of Taxa for Authors describing more than 100 species
One can see from examination of these figures that active work continues on the group. It is also evident from the graph above that a tremendous amount of work remains to be done just to form a basic outline of diversity in the group worldwide, especially in the Southern Hemisphere and particularly in Australia.
The subfamily and tribal classification presented in this database is consistent with, but does not necessarily reflect all aspects of, the most recently published literature. I would not argue that this is the only feasible system, but at least it is world-wide in scope. A thorough historical analysis of the use of higher taxon names will almost certainly require recourse to the Carvalho Catalog. A few cases deserve comment.
ISOMETOPINAE: This group was at one time recognized as a distinct family by most authors, and therefore not treated by Carvalho. Carayon (1958) and Schuh (1975) emphasized that Isometopinae share characters which are used to diagnose the Miridae; based on those arguments, the group is included here as a subfamily of Miridae. Herczek (1993) published a detailed tribal classification of the group. Version 2.2 of the catalog incorporates the classification of Herczek. Nonetheless, several new genera have been described by Akingbohungbe, Yasunaga, and others since the appearance of the work of Herczek (1993). Not all of these taxa area easily accommodated in the Herczek’s classification; I have treated some of them as incertae sedis and placed others in what seemed the most appropriate higher taxon. I agree with Yasunaga that the tribal classification of the group is in need of additional study, with the incorporation of a broader range of characters.
CYLAPINAE: The tribes of Cylapinae as recognized by Carvalho (1952) were almost certainly not monophyletic, and for that reason Schuh (1995) did not recognize them. This group has undergone extensive study since 1993. The higher classification has been re-evaluated and many new genera and species have been described. Version 2.2 of the catalog reflects the classification of the group largely as present in the subfamily-level catalog of Gorczyca (2006), the exception being that Vanniini is recognized as a valid tribe in this catalog. As with the Isometopinae, it would be my observation that the tribal classification of the Cylapinae is in need of additional study, with the incorporation of a broader range of characters and a more rigorous approach to understanding relationships.
PSALLOPINAE: This group, represented by a few described genera, has a checkered history of taxonomic placement and is here treated as a subfamily as it was in the work of Schuh (1995).
ORTHOTYLINAE: This moderately large grouping has been recognized by many modern authors. In the absence of a more sweeping phylogenetic framework, Schuh (1995) chose to recognize three tribes within the Orthotylinae: Halticini, Nichomachini, and Orthotylini. Some authors have accorded the Halticinae subfamily rank, but without strong arguments for this action. Several additional tribal groupings, all of which have characters usually used to diagnose the Orthotylini, have at times also been recognized; these include the currently recognized Austromirini and Ceratocapsini. The Ceratocapsini may have merit, but the taxon has generally been diagnosed on myrmecomorphic habitus, a very poorly defined attribute; furthermore, not all myrmecomorphic groups within the broader Orthotylini have always been included in the Ceratocapsini, as for example the exclusion of Sericophanes Reuter by Henry and Wheeler (1988). Many New World genera which fit the diagnosis have never been formally placed in the group. The Austromirini (see Cassis, 2008) will presumably include a very large number of presently undescribed taxa, both myrmecomorphic and otherwise. Recognition of Austromirini and Ceratocapsini at the tribal level creates a paraphyletic Orthotylini in my view. Nonetheless, I have chosen to include these two tribes and assign genera to them, as best as I can understand the published arguments for their placement, recognizing the need for further study of the residual Orthotylini and the need to divide it into monophyletic subgroups.
PHYLINAE: This group was badly commingled with the Orthotylinae in the classification of Carvalho, primarily because he placed genera in the two groups on the basis of an insufficiently detailed analysis of parempodial structure and did not believe that the male genitalia were diagnostic. Schuh (1974, 1976, 1984) made many changes in generic placement, not only within the tribes of Phylinae and Orthotylinae and their included tribes, but also with respect to other subfamilies, such as the placement of the Dicyphina in the Bryocorinae, rather than the Phylinae as done by Carvalho (1958). Many subgroupings of Phylini, in addition to those included in this database, have been recognized for the Palearctic fauna, but not applied on a broader geographic basis and are therefore not included here. Ongoing work will doubtless influence our views on the classification of the Phylinae, particularly with regard to the Phylini, an obviously paraphyletic grouping.
BRYOCORINAE: This taxon has a mixed history. Schuh (1976) rediagnosed the group and its included tribes. Whether or not the Bryocorinae is monophyletic is still in question, but at least the arguments for monophyly of the tribes are stronger than was previously the case. What seems certain is that Bryocorini of Carvalho was clearly polyphyletic. Palaucoris Carvalho was originally placed in a distinct subfamily. Whether or not it is best associated with the Eccritotarsina, as done here, is also open to question.
Generic assignments of species within the Dicyphina have been particularly confusing. The only work attempting a world treatment is that of Cassis (1986), available only through University Microfilms. The present database follows Cassis’s classification of the Dicyphina. Where new combinations were created by Cassis (1986), these are accompanied by catalog entries.
DERAEOCORINAE: This group is presented in substantially the same format as in the work of Carvalho. Although some groups, such as Deraeocoris Kirschbaum, may not be monophyletic, most of the tribes appear to be reasonably well defined, with the possible exception of the Termatophylini. The work of Cassis (1995) on the Termatophylini represents one of the only published efforts to test the validity of deraeocorine classification.
MIRINAE: This is the largest subdivision of Miridae. In the works of Carvalho (1959) and Schuh (1995) its classification appeared the least problematic of the larger subfamilies, in terms of recognition of the subfamily as monophyletic, the composition of the included tribes, and the conception of many of the genera. Although Schwartz (1987) showed that the tribes recognized by Carvalho were largely monophyletic, work published primarily since 1993 makes it abundantly clear that the genus-level taxonomy of the Mirini was stable primarily because few of the concepts had ever been put to rigorous test. This group has undergone extensive description, revision, and analysis in the last two decades, as the database reflects. Even though former garbage groups such as Lygus Hahn are now monophyletic, or nearly so, many taxonomic problems remain, two of which stand out. First, the immense Neotropical fauna is in need of rigorous revisionary work and better integration with the fauna of Ethiopian Africa. Second, the fauna of Australia remains poorly known, even though there is a large and diverse assemblage of Mirini on the continent.
REFERENCE TO ORIGINAL DESCRIPTIONS: Catalog entries are given for the original description of all available names of which I am aware in the Miridae. The list of names has been checked against the Catalogue of the Heteroptera of the Palaearctic Region (Kerzhner and Josifov, 1999) and the Catalog of North American Heteroptera (Henry and Wheeler, 1988) as well as lists of names published by Carvalho (Froeschner and Carvalho, 1987, 1990, 1995) and Wagner (Weber, 1976). Catalog entries are also provided for a relatively complete, although certainly not exhaustive, selection of morphological, biological, and faunistic references for all taxa mentioned in the literature during the years 1956 through 1993, with a lesser number of such entries from 1993 to the present. Unlike the Carvalho Catalog, this database does not include literature citations for taxon entries in catalogs, other than for the work of Carvalho, except for the few cases where there is new information on nomenclature or the status of names.
LITERATURE SEARCH: The literature included in this catalog was found through searches of the Zoological Record, Biological Abstracts, the new literature in the library of the American Museum of Natural History, and copies of papers sent by colleagues. Searching of the systematic literature was as thorough as practicable. The search of other literature has been motivated to find—in order of priority—host information, morphological information, non-control related biological information, and distributional data. Some references included in the bibliography have not been cataloged and are marked “NOT CATALOGED.”
ANNOTATIONS: Annotations of the references are similar to those in the Carvalho Catalog, but differ by indicating what type of figures are to be found, as for example, male and female genitalia, habitus illustrations, or scanning micrographs, and more consistently indicate which papers contain information on hosts. The following table contains abbreviations found in the annotations:
|biol.||life history information|
|FG||figures of female genitalia|
|MG||figures of male genitalia|
DISTRIBUTIONAL INFORMATION: A statement of distribution is associated with each reference containing geographic information based on the examination of specimens. The approach differs from that taken by Carvalho, in whose work distributional information was not directly associated with the reference from which it came. In the summary faunistic works of Wagner and other authors it is often not clear that the authors actually examined specimens in the preparation of their works; in such instances no distribution information is included in the catalog.
In many cases distributional information is recorded as found in the original reference, but in some cases it has been modified to reflect more modern political geography, or is based solely on geography. For example, taxa are often recorded in this catalog as being from Borneo rather than Kalimantan or West Malaysia, and from New Guinea rather than from Papua New Guinea or Indonesia: Irian Jaya; some records are listed as USSR, whereas most indications of USSR in the literature are listed in the catalog under the historical subdivision or modern state. Type localities are not listed as such. Where newly described species are recorded from a number of political or geographic entities, the one containing the type locality is usually listed first.
Coding of distributions by Biogeographic Region: Most records in the database that contain distributional information also receive a code that assigns them to a biogeographic region/subregion. This method of coding allows for more effective searching via the Geographic Distribution module. Such coding is, however, frequently subject to interpretation, where the dividing line between regions/subregions is not clear. Prominent examples include the distinction between the Oriental and Palearctic regions in China and the distinction between the Nearctic and Neotropical regions in southern Mexico and northern Central America.
HOST DATA: Host data are recorded for a significant number of taxa. These data are derived primarily from what I have interpreted to be primary observations. Because this catalog does not treat all of the literature on the Miridae, many mentions of host associations in the literature have not been captured. The following graph indicates the distribution of hosts by species. As can be seen, the majority of Miridae species for which hosts are known appear to breed on a single plant species.
Hosts are assigned to kingdom, family, genus, and species. Most host are plants; the taxonomic authority used to adjudicate family placement is The Plant Book by Mabberley.
TAXONOMIC HIERARCHY: The present catalog includes the following levels in the taxonomic hierarchy: family, subfamily, tribe, subtribe, genus, and species.
Subgenera: The exclusion of subgenera as valid taxa has forced abandonment of some of the hierarchic structure found in the literature, but in the final analysis was deemed the only feasible approach, for the following reasons. First, although subgenera have been frequently used by European authors, they are almost nonexistent in the taxonomy of the rest of the world fauna. This would present no problem if genera found in the Palearctic were restricted to that region, but such is not the case, and many species would have to be placed in the nominate subgenus, not by intent, but only because they have never been placed in any recognized subgenus. The very large, widely distributed genus Phytocoris Hahn, is an excellent example of the issue confronted with the recognition of subgenera. The use of subgenera further complicates a primary value of a catalog such as this, recognition of primary homonyms, which are based on the genus under which species are proposed, not the subgenus. Catalog entries include information on subgeneric placement even though the catalog does not group species by subgenus. All subgenera are treated as objective synonyms of their parent genera.
In most cases, the elimination of subgenera from the classification should cause little or no confusion. As of June 2011, the genus Orthotylus Fieber presents the most complex situation, my treatment of which may not satisfy all workers. Many species placed in Orthotylus probably have no close relationship to the European type species. Because the group has never been revised on a world basis, there is no diagnosis for the genus that can be readily applied on all continents. Many species — from the Holarctic, non-Ethiopian Africa, and the New World tropics — have been placed in Melanotrichus Reuter because they possess dark, flattened setae on the dorsum. Yet, even in the Palearctic, there is little agreement on which species belong to Melanotrichus, and whether or not this grouping should receive generic or subgeneric status. Thus, all species placed in Melanotrichus and several other segregates of Orthotylus appear under Orthotylus sensu lato. The exceptions are Brooksetta Kelton and Labopidea Uhler.
Subspecies: Subspecies are not recognized in the present work because there is little uniformity of application of this category, either taxic or geographic. Therefore, in addition to those species-group taxa that have been formally synonymized in the literature, all subspecies (and varieties and forms described before 1961 and not subsequently elevated to species) are treated as objective synonyms of their parent species.
FOSSIL TAXA: Carvalho listed fossil taxa at the end of his catalog. Although the number of such names was small, his approach made the discovery of homonyms much more difficult, except through the use of the index to his catalog. Fossil and Recent taxa are listed together in the present work. The number of described fossil taxa has increased dramatically in the last few decades, most of these being from amber.
GENDER AGREEMENT IN SCIENTIFIC NAMES: The inability of most authors--both past and present--to deal with classical languages has produced problems with gender agreement in scientific names. Many authors have gone to lengths to avoid the problem, by using geographic names and nouns in apposition. Some entomologists, notably the late George Steyskal, have taken it upon themselves to correct the nomenclature as published in catalogs of the Heteroptera (Steyskal, 1973). In some cases there would seem to be little disagreement about Steykal’s arguments for changing the endings of names. A good example might be the species of Stenodema Laporte included in the Carvalho catalog. Other groups seem to present a different situation, however. For example, Campylomma Reuter was treated as feminine by its author, as can be seen from the termination of the originally included species. Yet, Steyskal (1973) argued that the generic name should be treated as neuter because the grammar aids in the code indicated that all names ending in -omma be treated as such, and therefore, not only is the gender of the type species incorrect, but also the gender of nearly all subsequently described species is incorrect. The present catalog corrects obvious conflicts and inconsistencies, but does not modify the gender selection of the majority of classical authors who wrote descriptions in Latin. This approach is not in all cases in agreement with that of Kerzhner and Josifov (1999) for the Palearctic fauna. As a general rule, I have not “corrected” the literature, but simply recorded what I saw.
ORIGINAL AND CORRECT SPELLINGS: Some names were clearly printed in error in the original publication, as for example the misspelling of the name of the person being honored in the creation of a patronymic, or multiple spellings of a name in a single publication. The current catalog adopts the spelling which was most obviously intended, although it may not be the one used in the main reference to the taxon. In some cases the original spelling is retained although subsequent authors may have adopted variants. Some names appear to be incorrectly formed, but are certainly valid under the code and I have therefore retained them in most cases, even though they are orthographically incorrect in Latin. An example is the use of the term nigrus by Carvalho for species that are black.
UNAVAILABLE NAMES: As noted above, the catalog includes all available names of which I have become aware. It does not list names which were first published as unavailable. Neither does it contain references to available names which, after 1960, are used for categories which as of that time are not subject to the rules of availability.
CONFIRMING THE CONTENTS: To the degree possible, every entry in the catalog has been checked against original literature. This has not been possible for a relatively small proportion of references, which are marked “NOT SEEN” in the bibliography.
Many persons participated in the preparation of this catalog and its underlying database, and without their help it might never have been completed. They are listed by category of activity.
First, I thank the late Jose C. M. Carvalho for his blessing to produce a supplement to his singular work, and for his many words of encouragement. Without his catalog, the taxonomy of the Miridae would certainly not have advanced to the state in which we find it today, and almost certainly many more homonyms, synonyms, and taxonomic misplacements would have been published. In every sense, Carvalho deserves recognition for his seminal contributions to modern-day mirid classification, as did O.M. Reuter before him and F.X. Fieber before him. I can only hope that Jose Candido, as he was know to his friends, would have found the print and on-line versions of the Plant Bugs of the World catalog worthy successors to his own.
Most of the bibliography through 1992 was assembled, prepared in computer form, and checked for accuracy by Brenda Massie. Allma Edwards typed the literature section of the Carvalho catalog from which the database bibliography for the literature prior to 1956 was generated. The late Leslie Marcus assisted with the original computerization of the bibliography.
Library assistance was provided by Sule Oygur Fischl, Paula Miller, Ruth Salas and the library staff of the American Museum of Natural History. Thomas J. Henry and Richard Froeschner assisted with literature searches at the Smithsonian Institution. The late I.M. Kerzhner provided some difficult-to-find references dealing with the Palearctic fauna. I thank those colleagues who have faithfully sent reprints of their works over the years. The effort they expended was more than repaid by the benefits provided in the preparation of this catalog.
A very large proportion of the catalog entries pertaining to the literature published between 1956 and 1985 was prepared by Bella Galil. To her I owe a great debt that cannot easily be repaid in words.
Checking of information in the catalog through about 1998, including the bibliography, was greatly facilitated by I.M. Kerzhner and his work on the Palearctic fauna. Further checking was done against the carefully prepared Miridae chapter of the Catalog of Heteroptera of America North of Mexico (Henry and Wheeler, 1988). I, however, take complete responsibility for the final form of this catalog and the errors contained in it. I am under no illusion regarding errors, for to produce a catalog without errors would be to produce no catalog at all.
Entry of much of the data into the database through 1990 was done by Bea Brewster and Gail Motyka. Their ability as typists and dedication to the sometimes tedious task helped form the core of the catalog.
For creation of the original Wang database application I am grateful to Joan Whelan and Gail Motyka. Through their cooperative efforts we developed what would now be regarded as a primitive relational model for acceptance of data collected in the first phases of the project.
The relational database application upon which the 1995 printed catalog was based was written by Gary M. Shapiro, Brooklyn, New York. The basic elements of the relational model were originally suggested by Robert Raven, Queensland Museum, Brisbane, Australia. The web interface and associated queries for on-line Versions 1, 2, and 2.1 were written by Mark Breedlove, Department of Library Services, American Museum of Natural History. Mark also designed the database schema and prepared the web implementation for Versions 2 and 2.1. Data entry for version 2.2 was made possible through a web-based application written by Simon Chang, Octeva Consulting, LLC. Ryan Choi of the American Museum of Natural History provided assistance with the database and the data pertaining to version 2.2.
Lee Herman and F. Christian Thompson contributed to the conception of the Schuh catalog, and spent many hours discussing the niceties of nomenclature, and database applications for hierarchic systematic problems in the case of Thompson. I also thank the late James A. Slater for his interest in this project and his many helpful and encouraging suggestions.
Members of the NSF-funded Planetary Biodiversity Inventory for the study of the subfamilies Orthotylinae and Phylinae used the database extensively and made many suggestions for its improvement. These include, but are not restricted to Michael D. Schwartz, Dimitri Forero, Dinise Wyniger, and Christiane Weirauch.
The National Science Foundation made possible the preparation of the database application upon which the published catalog of Schuh was based and generously supported the publication of the finished work. Additional support came from a grant from the Global Biodiversity Information Facility (GBIF) and from the Planetary Biodiversity Inventory Award from the NSF.
This work is dedicated to the memory of the late Dr. Thomas D. Nicholson, former director of the American Museum of Natural History. It was his appreciation of value of works such as this, and his willingness to provide financial support in the initial stages, that made this project possible. I only regret that he did not live to see the finished printed or on-line products.
— Randall T. Schuh
Citations for this site should read: “Schuh, R.T. 2002-2013. On-line Systematic Catalog of Plant Bugs (Insecta: Heteroptera: Miridae). http://research.amnh.org/pbi/catalog/”
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