Version 2 of the Plant Bugs of the World (Insecta: Heteroptera: Miridae) on-line database is based on a revised MySQL schema and offers a more flexible approach to querying the data than was available in Version 1. Information can be found using the search boxes or by browsing the taxonomic hierarchy.
Browse the taxonomic hierarchy:
Search boxes allow for two searching strategies:
Scientific name searches:
Host searches:
Bibliographic searches:
Hierachic 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. 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:
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 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 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 pages can now be viewed through the digital library for the subfamilies Orthotylinae and Phylinae. Corrections to the catalog published by Kerzhner and Schuh (2001) are included in the database, as are most new names and other nomenclatural changes published between 1993 and 2001. Most entries are based on direct examination of the primary literature. A few entries are taken directly from the work of Kerzhner and Josifov (1999) without further verification, because copies of the original literature were not available.
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 intervening 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 in the taxonomy of the Miridae is nonetheless remarkable. The following tables provide data on historical growth in our knowledge of the Miridae.
Table 1. Numbers of Available Names
|
Genus-group names |
|
|
Described before 1956 |
1,145 |
|
Described between 1955 and 1994 |
678 |
|
Described after 1993 |
120 |
|
Available |
1,942 |
|
Valid |
1,507 |
|
Species-group names |
|
|
Described before 1956 |
6,637 |
|
Described between 1955 and 1994 |
5,465 |
|
Described after 1993 |
945 |
|
Available |
13,048 |
|
Valid |
10,040 |
Table 2. Numbers of Papers Published
|
Cataloged papers published before 1956 |
1307 |
|
Papers cataloged published between 1956-1993 |
1927 |
|
Papers cataloged published after 1993 |
354 |
|
Total |
3634 |
Table 3. Numbers of species described, by decade.
|
1758--1767 |
40 |
|
1768--1777 |
24 |
|
1778--1787 |
69 |
|
1788--1797 |
73 |
|
1798--1807 |
99 |
|
1808--1817 |
2 |
|
1818--1827 |
11 |
|
1828--1837 |
144 |
|
1838--1848 |
93 |
|
1848--1857 |
197 |
|
1858--1867 |
342 |
|
1868--1877 |
396 |
|
1878--1887 |
568 |
|
1888--1897 |
467 |
|
1898--1907 |
714 |
|
1908--1917 |
1304 |
|
1918--1927 |
735 |
|
1928--1937 |
477 |
|
1938--1947 |
385 |
|
1948--1957 |
695 |
|
1958--1967 |
904 |
|
1968--1977 |
1804 |
|
1978--1987 |
1480 |
|
1988-1997 |
1679 |
|
1998-to date |
523 |
Table 4. Numbers of Taxa for Authors describing more than 100 species
|
Author |
Generic-group |
Species-group |
|
Carvalho |
393 |
2153 |
|
Knight |
45 |
1345 |
|
Reuter |
260 |
1122 |
|
Poppius |
181 |
735 |
|
Linnavuori |
43 |
729 |
|
Wagner |
73 |
544 |
|
Distant |
154 |
515 |
|
Schuh |
66 |
421 |
|
Stichel |
4 |
312 |
|
Van Duzee |
16 |
235 |
|
Stonedahl |
9 |
218 |
|
Schaffner |
35 |
181 |
|
Ohdiambo |
27 |
163 |
|
Stal |
17 |
163 |
|
Uhler |
29 |
154 |
|
Fieber |
102 |
141 |
|
Kerzhner |
32 |
189 |
|
Kelton |
8 |
121 |
|
Henry |
14 |
120 |
|
Lindberg |
8 |
112 |
|
Horvath |
9 |
104 |
|
Kirkaldy |
71 |
101 |
One can see from examination of these figures that active work continues on the group. It is also evident that a tremendous amount of work remains to be done just to form a basic outline of diversity in the group worldwide. At present the most glaring hole is the absence of published information on the Australian fauna.
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 Isometopine 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. The many tribal-level divisions included in Herczek"s work are at present not incorporated into the database.
CYLAPINAE: The tribes of Cylapinae as recognized by Carvalho (1952) are almost certainly not monophyletic, and for that reason Schuh 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. At present, the higher classification in database appears as it did in the work of Schuh, although references to newly described higher taxa are given.
PSALLOPINAE: This group, represented by a few described genus, has a checkered history of subfamily placement and is here treated as a subfamily as it was in the work of Schuh.
ORTHOTYLINAE: This moderately large grouping has been recognized by many modern authors on the basis of pretarsal structure. However, the Pilophorini (Phylinae) have parempodia that are apparently identical with those of the Orthotylinae, and therefore these structures may not justify the monophyly of the group. The male and female genitalia, furthermore, show substantial variation, and appear to offer little argument for the monophyly of the group. In the absence of a more sweeping phylogenetic framework, Schuh (1995) to recognize three tribes within the Orthotylinae. Some authors have accorded the Halticinae subfamily rank. Several additional tribal groupings, all of which have characters usually used to diagnose the Orthotylini, have also been recognized, but are not included in the present work. These include the Ceratocapsini. This group may have merit, but it 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. Recognition of these groups at the tribal level unqestionably creates a paraphyletic Orthotylini. Further work will be required before a natural classification is available.
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 alone were diagnostic. Schuh (1974, 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. 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.
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 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 subfamily and tribal placement for genera formerly placed in the Dicyphini by Carvalho, as well as Cassis assignment of species to genera. Where new combinations were created by Cassis (1986), these are accompanied by catalog entries.
DERAEOCORINAE: This group in presented in substantially the same format as Carvalho. Certainly some groups, such as Deraeocoris Kirschbaum, are not monophyletic, although most of the tribes appear to be reasonably well defined. 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 generic-level taxonomy of the Mirini was stable only because few of the concepts had ever been put to rigorous test. This group has undergone extensive analysis in the last decade, as the database reflects.
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 published for 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 of taxa 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:
|
combination |
|
|
gen. |
genus |
|
n. |
new |
|
subgen. |
subgenus |
|
sp. |
species (singular) |
|
spp. |
species (plural) |
|
ssp. |
subspecies |
|
syn. |
synonymy |
|
biol. |
life history information |
|
descr. |
description |
|
diag. |
diagnosis |
|
disc. |
discussion |
|
dist. |
distribution |
|
DV |
habitus illustration |
|
FG |
figures of female genitalia |
|
hab. |
habitat |
|
host |
host |
|
LV |
lateral view |
|
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, whose 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 that work; 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 from Borneo rather than Kalimantan, 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 usally listed first.
TAXONOMIC HIERARCHY: The present catalog includes the following levels in the taxonomic hierarchy: family, subfamily, tribe, subtribe, genus, and species. This approach 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 taxa 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 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 contain subgenera.
In most cases, the elimination of subgenera from the classification should cause little or no confusion. The genus Orthotylus Fieber presents the most complex situation, the 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 a world basis. 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.
Neither are subspecies 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 synonyms.
FOSSIL TAXA: Carvalho listed fossil taxa at the end of his catalog. Although the number of such names is small, his approach made the discovery of homonyms much more difficult, except through the use of the index. Fossil and recent taxa are list together I the present work.
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 indicate 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.
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 all cases, even though 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 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 and synonyms would have been published. In every sense, Carvalho must be considered the father of modern-day mirid classification, as was 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 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, 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. 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 were prepared by Bella Galil. To her I owe a great debt that cannot easily be repaid in words.
Checking of information in the catalog, 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), whose authors are to be commended for their efforts. I, however, take complete responsibility for the final form of the catalog and the errors contained. 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 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 a basic relational model for acceptance data collected in the first phases of the project.
The relational database application upon which the 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. The web interface and associated queries for on-line Version 1 were written by Mark Breedlove, Department of Library Services, American Museum of Natural History. Mark designed the database schema and prepared the web implementation for Version 2.
Lee Herman and F. Christian Thompson contributed immeasurably to the conception of the Schuh catalog, and spent countless hours discussing the niceties of nomenclature, and database applications for hierarchic systematic problems in the case of Thompson. I also thank James A. Slater for his interest in this project and his many helpful and encouraging suggestions.
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
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