Table of contents
The Agromyzidae are known primarily as leaf-miners but a significant number of species are internal stem-borers and seed feeders and, to a lesser extent, root or bud feeders and gall causers. Monocots, dicots and horsetails are attacked and the thalli of liver worts and horn worts are also known as larval substrates. Larvae of the genus Phytobia are only known to feed on the youngest xylem inside the cambium zone of trees belonging into many plant families but their "pith flecks" or medullary spots in the timber - cross-sections through the feeding tracks of the larvae (26700:1933) - are only confirmed for two conifer genera, Callitris in Australia and Juniperus in Greece. Without any exception, the larvae are true phytophagous plant parasites. No fungus feeder is known. In adaption to this development, the asymmetric larval mouth hooks are singular among the Diptera Cyclorrhapha, serving as a scythe-like mowing tool of the maggots.
As a second peculiarity, without exception the larvae are moving alternatively on their left or right body side within the living plant tissue, which enables them to scrape out plant cells and chloroplasts. The complicated way of feeding is demonstrated for a larva of Phytomyza ilicis (3673:Taf. 53). The plant pulp is sucked in into an asymmetric mouth slit between the right and left base of the mandibles (figures 6297:311). As a result of this peculiar morphology without any counterpart in fly larvae, the frass particles are deposited not in a straight central line but alternatively along the right and left edges inside the leaf mine. Several frass pellets are deposited at one side of the mine, then the larva rotates by 180° and consequently the frass is now deposited at the opposite side of the mine during. The larva is now moving on the other side of its body. Thus, leaf mines of Agromyzidae can be easily distinguished from all other leaf mining insect larvae. Only those mines produced by subcuticular epidermal leaf-mining species (for example those belonging into the genera Tropicomyia or Kleinschmidtimyia) are more difficult to assign to agromyzids as such species only ingest cell sap from the chloroplast-free epidermis cells. Consequently, they produce only greenish "clouds" of excrement or no frass at all. In doubtful cases, the remaining typical mouth hooks of the first and 2nd instar inside the mine let identify the agromyzid mine causer, as well dead third larval instars or puparia.
A further exception from other fly families is the specific drilling mechanism hidden within the black and conical oviscape (oviscapt, ovipositor sheath) at the hind end of the female flies (Fig. 6217:3; 26700:1913,1929; 6217:3). Equipped with a multitude of black teeth being arranged in a special pattern, the female agromyzids drill small "feeding punctures" (keyword: feeding punctures) into the leaf or stem surface of their host plants or other plants. Exactly 200 were drilled during one session observed in a leaflet of Golden Chain (Laburnum anagyroides) being produced by Agromyza demeijerei (6298:Tafel 14). After each drilling procedure, the female steps backwards and ingests the tiny protruding droplet of plant sap. For egg deposition the drilling apparatus is also used, but the drilled epidermal hole is more oblique and gets the shape of a small pocket in which the single egg is laid. The omnipresent ants in tropical forests gnaw away the drilled epidermis and pull out the egg, one reason for the rareness of agromyzids in the tropics. Normally, only one or few eggs are laid per leaf, which prevents later larval competition. Interesting exceptions occur, for example Agromyza megalopsis always deposits several eggs in crossline into a cereal leaf producing racing siblings (6298:Tafel 13). Finally, such a group excavate the whole part of the leaf downwards of the group of their emty eggshells. An extreme example is Cerodontha iridis: groups of about hundred larvae at close quarters pupariate in one or two long row(s) within one Iris leaf (M.v.T., Oxford Univ. Park).
A multitude of works for the identification of families of Diptera are included in "World Bibliography of Agromyzidae and Chloropidae Online" and more than 100 references are equipped with the keyword "family keys". The ten most actual and complete ones since 1973 can be entered using the BIBLIO numbers 25669, 21807, 25019, 21343, 22060, 17626, 8682, 10169, 15961, 10202 (sequence starts with latest year and fantastic work available free of charge online at https://www.nhm.ac.uk/our-science/our-work/biodiversity/manual-afrotropical-diptera.html). In German language and focused on Central Europe, the BIBLIO numbers 20899, 22514 and 28111 may be added, later editions are available.
Agromyzidae (leaf-miner flies) belong into the so-called Acalyptratae Macquart, 1835 (often misspelled as Acalyptrata). This group of "true" flies include 61 families (23354 und Clusiomitidae), most of which can be grouped into superfamilies. To date, for Agromyzidae a phylogenetic sister group cannot be identified and the long time used superfamily Opomyzoidea was rejected for them (26700:1022). Acalyptates normally (exception for example Platystomatidae, Psilidae) lack a dorsolateral longitudinal V-shaped seam in their 2nd antennal segment, the 'pedicel', they do not possess a large thoracic squama (landing flap) below the wing base, their dorsal thoracic transverse suture is widely interrupted in the anterior centre of the scutum (=mesonotum), their single outer postalar seta is not inserting on a small callus being surrounded by a weak groove, and tibiae miss long strong bristles or they possess only one or few short setae. All those features apply for world Agromyzidae.
Shared with many other acalyptrates and the characteristics used in all identification keys are the following morphological features: Size 0.5-6.0mm; wing always with only one costal break near the end of subcosta; subcosta fading away distally or melting with vein R1; vein R4+5 and vein M1 distally never convergent; postverticals (=postocellars) divergent; vibrissae present [sometimes vestigial, for example Ophiomyia anomala, Selachops flavocinctus, Phytomyza gymnostoma]; tibiae always without a dorsal preapical seta.
All Agromyzidae possess parafacialia (cheeks) running from the eye margin at level of antennal base downwards along face and continued along anterior and whole lower eye margin. They are well separated from the ventrally adjoining gena (=jowl) and sometimes they are widened or slightly swollen. In agromyzid literature, in the relation of eye depth (=eye height) compared to depth of gena the parafacial strip is included and both together - cheek plus jowl - are incorrectly addressed as jowl or gena only. In two published photos (25604:39,45) the cheeks are well documented, an extremely deep cheek is figured for Liriomyza lutea (25584:165). The literature on identification of families of Acalyptratae only exceptionally mentions the presence or absence of parafacialia.
The females of all Agromyzidae possess a conical sclerotized oviscape being a closed sheath around the complicate retracted ovipositor and the hidden ventral Receptaculum seminis). The oviscape is never dorsocentrally flattened like in several families of the Tephritoidea (for example Platystomatidae, Lonchaeidae) and only very seldom it is slightly compressed laterally (Liriomyza taurica). The Australasian/Indian family Fergusonididae possess a similar looking oviscape, as well Pseudopomyzidae and most Tephritidae. The female cerci are bearing several soft and partly stiff setulae [an exeption is Gymnophytomyza heteroneura with one tiny peg on each long cercus] and the cerci are not fused and sclerotized as a piercing mechanism. The ovipositor of all Agromyzidae is covered with a multitude of denticles arranged in a pattern of many oblique rows, used for drilling holes into the alive plant tissue. Only two sclerotized spermathecae are present.
Female and male Agromyzidae can be identified as follows: the frons of males and females is always of the same width, and also other sexual dimorphic differences known from "true flies" play no role for family assignment of Agromyzidae; one or more upper orbital setae (ors) are present [only three exceptions without ors are Selachops flavocinctus and males of Ophiomyia pinguis and O. nasuta], which are reclinate or additionally slightly bent outwards, the posterior one of two ors may be shorter; one or several lower (=anterior) orbitals (ori) always are strongly curved inwards [one exception in the high Andes above 3000m a.s.l.: male of Cerodontha nigricornis without ori]; inner and outer verticals present [very few exceptions]; postverticals (pvt, =postocellars) always divergent [absent in Ophiomyia arabica, Xeniomyza ilicitensis]; ocelli and divergent ocellar setae (oc) present; 1 supracoxal propleural (pp) seta present [absent in Gymnophytomyza heteroneura and at least one Penetagromyza species]; peristomal setae and at least 1 anepisternal (m, =mesopleural) [the latter absent in Xeniomyza intermedia and Selachops flavocinctus]; at least 1 katepisternal (st, =sternopleural) present; 1 humeral (h), 1 presutural intra-alar (prs) and 1 exterior postalar (epa) always present; 1 supraalar seta (sa) present [one exception is a Penetagromyza sp. n. from South Africa with absent sa]; 2 notopleurals (n) present [1 n only in some species of subgenus Cerodontha and in Cerodontha (Poemyza) sudeshi, as well in few Phytomyza and Ptochomyza spp.]; 1 basal/lateral scutellar seta (bs) present [absent only in subgenera Cerodontha and Xenophytomyza, vestigial in Oriental Cerodontha (Poemyza) oriziphila and in some species belonging into Cerodontha bisetiorbita and C. setaria species groups (20127:111)]; 1 apical scutellar (as) present [one exception is ?Amauromyza sp. n. from 4,200m altitude in the high NE Andes of Chile without this bristle, a case being very rare in Schizophora]; 2 up to 7 dorsocentrals (dc) present [one exception in northern Iran: Agromyza sp. n. with 1 dc only]. The following setae belonging to the ground pattern of the family may be present or absent: acrostichals (acr) being always arranged in rows; orbital setulae; 1 pair of prescutellars (prsc); 1 postsutural intra-alar (ia); 1 inner postalar (ipa); 1-3 (rarely more) small posterolateral (=posterodorsal) fore- and mid-tibial setae; scutellum and frons are bare but some tiny hairs may rarely occur on scutellum [one exception: Selachops flavocinctus with haired scutellum]; rarely some tiny hairs along frontal edge of frons, for example Melanagromyza caerulea and large Agromyza and Phytobia spp.; arista always present, appearing bare, pubescent or plumose [exception: Japanagromyza sp. n. from Peruvian rain forest with 25 long rays on arista (6207:303)]; wings neither absent nor vestigial; eyes without strong metallic sheen and without differently coloured transverse fascia; shape and size of eye normal [exception one Ophiomyia sp. n. from Petra (Jordan) with downsized eyes and very large gena]; pictured wings are rare and occur in the tropics (26881:Fig.1); anal cell and shortened anal vein present [some few species without anal vein, for example Amauromyza obscura]. Larvae move on their lateral sides, their basally combined mouth hooks are always asymmetric and always working like mowing with a scythe inside the live plant tissue (video see 28000: https:/agromyzidae.co.uk/feeding-habits).
In "World Bibliography of Agromyzidae and Chloropidae Online", for about 60 references the keyword "keys to genera" is added. The seven most recent detailed keys in English language are found in 26700, 27333, 23929, 22123, 8851, 8869, 8825. Earlier sources no longer agree with the actual definitions of certain genera. No key exists for all valid taxa of the genus group.
The index of the world Agromyzidae includes about 5,401 scientific names with 9,523 file cards with information on names and their source. The collection of literature was started in 1966 and continued until the end of 2022. Some few sources and new taxa published between January 1 and May 31 were included, too. An EXCEL list with the information in the header of each card has been maintained since the use of a computer in 1998. In addition to the scientific name are following: author(s), year and - if necessary - round brackets around both, finally after a dot genus and in round brackets subgenus. If a name actually is invalid (being a junior synonym, nomen dubium, nomen oblitum, nomen nudum, misspelling or another error) the complete name is set in square brackets. All found published phylogenetical clade names (being outside the Rules of the ICZN) are collected for the first time and are included in the list, too. They are underlined in red colour, are combined with the word "group" or [German] "Gruppe" and are positioned before the valid or invalid species or genus name. Thus, their position is the only exception of the strict alphabetic order of all names. The alphabetic sequence was not automaticly generated by EXCEL but thoroughly by eye: All words and year-numbers in a name have been used as if they are combined to one single long word. For this ordering only brackets have been neglected. Family group names ending with …idae are only added to a header name if that one actually belongs into another family of Diptera or to a homonymous other animal. Only in the header of the first of several cards for a name all further genera and subgenera in which the species ever had been treated or listed are noted in square brackets. Multiple cards (up to 97 for Liriomyza trifolii) exist for many names. They are numbered consecutively by an encircled number at the top right. Just below this number partly a date is added (day, month, year). It indicates the day when this further card was established with its first entry.
Nevertheless, if it is doubtless that a species was misidentified in a used source, the reference is listed on the card with the erroneously used name. There are some few exceptions for such cases when the correct actual valid name could be found out for the misidentification. In such exceptions, the citation has also an entry on the card for the correct name but this is set in brackets. Phytomyza rufipes is such an example.
Below the header of a card further information follows: All found publications relating to the name, abbreviated as Author(s), year, exclusively all exact pages in the article or book, partly notes on the contents or taxonomy and in many cases even the BIBLIO number of the database at the right end of a line. Red underlined notes in certain references apply to the original description or first publication of an actual taxonomical status in the year 2022. Lower case letters behind the year were not used for multiple publications of an author within one year. In most such cases the given pages exclude errors in selecting the belonging article or book in "World Bibliography of Agromyzidae and Chloropidae Online". In early written entries for some few authors [for example Hering; Spencer] such lower case letters were used. As not repeated in "World Bibliography of Agromyzidae and Chloropidae Online" they should be neglected.
In addition, there are cards with further information that can be found using "further information". They are restricted to generic names. Those lines appear alphabetically after the genus in question. An example is the sequence Melanagromyza spec., M. spec., M. spec. nov., M. spp. [the latter meaning several unidentified species], M. with atypical aedoeagus [the German and Latin term for aedeagus], M. with plumy arista, etc., altogether 18, 11, 1, 3, and 25 cards, respectively. The first 33 cards refer to published notes, the 25 latter ones are subjectively selected by the author for morphologically interesting details which could be used for the construction of keys or for phylogenetical and functional-morphologic arguments. Of course, the latter 25 are incomplete concerning the morphological keywords and as well for the number of listed world species for each [yellow] highlighted keyword. Here must be said that all common references to the exemplary taxon Melanagromyza can be found only on the first 18 cards for Melanagromyza Hendel, 1920.
The header of the file cards corresponds to the information in the EXCEL list and the data set in the database. All names, valid or not, species group, genus group, family group and nomenclaturally not protected clade names are presented in one single list. In order to publish the world index of Agromyzidae on the Internet, the file cards and the basic EXCEL list were thoroughly revised which required several years labour. The data from the EXCEL list were transferred to database tables. They were initially used for indexing all file card images and now used for accessing the images by name, author, year, etc. All file cards were scanned exactly according to their order in the file card boxes. If the backs are also used, the images follow the respective front. Processing using OCR (optical character recognition) was not planned.
Via the menu ☰ at the top edge of the screen you can reach the settings, this introduction, M. von Tschirnhaus' bibliography and the database "World Chloropidae Online" as well as further links to Diptera. Under the menu item Settings you can change the language of the controls and some data in the hit list, for example in the columns "Rank" or "Valid". Choose German or English. Enter the number of rows that you want the hit list to display and then click "save". A cookie on your device will now contain this information for future visits to the website. If you choose "do not save", no cookie will be saved and the program will use English and 10 lines.
Below the menu the scientific names (=rows of the table, =data records) are displayed in the form of a table. At startup, this hit list contains all names in undefined order. Only as many rows are displayed as defined in Settings. However, only the 8 columns that can be searched are visible. All further information can be found under "World Agromyzidae Online details". To do this, click on the data sheet icon to the left of each scientific name. Finally, the number in the blue circle signals the number of file cards associated with the name.
The pagination buttons are located below the hit list. Depending on the position in the database, you can scroll back with the left and forward with the right buttons. The page number and the count of the visible data records are given between the buttons.
Searching for the name you want and sorting the lines of the hit list is done using the controls in the three-line table header. A selection of sorting operators is hidden below the column title , a selection of search operators below the second header line , and the search term can be entered in the third line . Active operators and search words are displayed in blue font on a light background. The question in the example above can be read from top to bottom as: Search for lines from Agromyzidae for which: scientific name starts with abb .
The left column of the table, called "Action", shows two buttons in the table header. With the button submit, the previously entered query is sent to the server. The button reset triggers a defined initial state, theat is all scientific names are included in the list, sorted alphabetically and the first lines are displayed.
For each line in the table body, the action column shows the data sheet icon with the number of associated file cards. Clicking on it will take you to the details page.
The last line of the hit list contains the button for downloading the selected data. Depending on the settings in the web browser, they are displayed immediately or stored in the download directory.
The file cards belonging to the scientific name are displayed on the left. If there is more than one card, the pagination buttons can be used to scroll through the deck of cards.
Below this (usually on the last file card) there are annotations that were not listed on the file cards.
On the right you will find further data, such as original citation with page number, comments, etc.
"Find references by" (bottom right) creates a link to "World Bibliography of Agromyzidae and Chloropidae Online". From the citations on the card, the first author and the year can be entered here, separated by space. Often the number from the BIBLIO database is also available. It can be used alternatively. After pressing the enter key or clicking on the symbol of the magnifying glass, the citation is retrieved from the associated M. von Tschirnhaus' "World Bibliography of Agromyzidae and Chloropidae Online" and displayed.
Finally, you can return back to the hit list with the button back to hit list at the bottom right. Please do not use the "back" button in your web browser!
At the beginning or after clicking on reset there all scientific names are included in the hit list. To search the records based on values in the columns, the search method and the value can be typed in rows 2 and 3 of the table header.
The search method is initially set to the most commonly one used. Click on the word to change it. In the input list that now appears, select an operator, for example "starts with".
To enter a search term, click on the input field, for example below Author, and enter the name you're looking for. Then click submit. All columns selected for searching, the search operators and the search words entered are displayed in blue font on a light background.
The table below explains how to search in the various columns.
| Column | Method | Explanation |
|---|---|---|
| Name | starts with | Finds scientific names beginning with the entered search term. |
| is equal to | Finds scientific names equal to the entered search term. | |
| contains | Finds scientific names containing the entered search term. | |
| Author | is equal to | The search for authors who have published the scientific name is made easier by a list of the different spellings of the personal names. Searching with Rodendorf leads to scientific names of Rodendorf, Rodendorf-Golmanova and Rohdendorf-Holmanová. |
| starts with | The operator gives the best result when the end of a name is uncertain. The truncation character "%" does not have to be entered at the end of a word. | |
| contains | The operator gives the best result when the spelling of a name or the position in the author's team is uncertain. The truncation character "%" does not have to be entered at the beginning or at the end of the word. | |
| Year | is before | Finds the scientific names published before the entered year. |
| is equal | Finds the scientific names published in the entered year. With Year is equal [to] 0 you will find data records for which no year could be determined. For some scientific names, a year is meaningless, the column is empty. | |
| is after | Finds the scientific names published after the entered year. | |
| Rank / Valid | is equal to | The search words are given in the associated selection list. To open this list, click on the caret pointing down in the input field. "Valid" is to be understood in the Agromyzidae system. |
| Genus | The search using genus names works in the same way as column "Scientific name". | |
| Region | contains | The search is done with the so-called Boolean full text search. The operators + (plus) and - (minus) indicate that a word must be present or must not be present in order for there to be a match. Several words separated by ; (semicolon) can be entered in the input field below the title of a column. Truncation with % (percent) is possible at the end of words. Finally "" (double quotation marks) are used to enclose a fixed term that can consist of several words. Zoogeographical regions are stored with "Afrotropic", "Australasian", "Nearctic", "Neotropic", "Oriental" and "Palaearctic". The region mentioned in the original description is marked with "²". For example, entering Palaearctic² (without space) will find all names that were first described for this region. In contrast, entering Pal% will give you all names for which the Palaearctis region has been mentioned in the literature (both "Palaearctic" and "Palaearctic²"). For some names, several regions have been published. To exclude all others except Palaearctic, enter +Pal%; -Afr%; -Aus%; -Nea%; -Neo%; -Ori% (+ and - and words separated by ;). |
| More info | contains | The column "More info" combines information from the EXCEL table from the columns "Original genus", "Other genera", "Other citations",and "Valid oldest name", and "State" for searching. In the detailed view, these data are assigned to the original columns of the EXCEL table. The Boolean full text search is carried out as described under "Region". |
| Original genus | Contains the original genus name. | |
| Other genera | Contains further genus names. | |
| Other citations | Contains an abbreviation and the corresponding BIBLIO number together with the page number. The most important abbreviations are: coll. = name in a collection, comb. = new combination, litt. = name in the literature, nom. = new name, ref. = reference to a source, stat. = revised state, and syn. = synonym. Other remarks are self-explanatory. The named BIBLIO number can be entered next to "Find reference by" and the citation immediately displayed. | |
| Valid oldest name | Contains the valid oldest name. For example, if you are looking for all synonyms of "Phytomyza ranunculi (Schrank, 1803)" enter ranunculi below "More info". You will get 29 names: albipes Meigen, 1830; bicolor Anonymus, 1848; cinereo-vittata Staeger; ...; zetterstedtii Schiner, 1864. The hit list also contains 2 valid species (caulinaris Hering, 1949 and stolonigena Hering, 1949) and one group name (notata Meigen, 1830 group). To exclude these, additionally select species group name below "Rank" and no below "Valid". |
|
| State | Contains the original classification of the name, partly processed in the columns "Rank" and "Valid": error in the widest sense, fossil, family transfer, homonymous genus in zoology, invalid family group name, invalid genus group name, invalid subgenus combination, nomen dubium, nomen nudum, nomen oblitum, preoccupied, species in a homonymous genus, valid subspecies, synonym incl. variety, teratological, unjustified emendation, unavailable name, valid family group name, valid genus & subgenus, valid species, valid subgenus combination, genus clade, species clade, junior species clade name For example, if you are looking for names of fossil species, enter fossil below "More info". |
After clicking reset, the lines in the list of hits sorted by scientific name. The current sorting can be recognized by the blue font on a light background and the arrow in front of the column title in the table header. In addition, a small superscript number indicates the order in which the columns were sorted.
The column titles are used to change the order. To do this, click on it, for example on Author. In the input list that now appears, select ↓descending (Z to A), unsorted or ↑ascending (A to Z) and click on submit.
In general, the scientific names are sorted alphabetically in ascending order. If there are several lines with the same name, groups come first and additional information last.
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Please cite the database as follows: von Tschirnhaus, M. & Groll, E. 2024: World Agromyzidae Online. - https://sdei.senckenberg.de/tschirnhaus-agromyzidae
Vignette: Ptochomyza asparagivora, Photo André Burgers