Pest Fruit Flies of the World - Larvae

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L.E. Carroll, A.L. Norrbom, M.J. Dallwitz, and F.C. Thompson

Character List

Nomenclature #1. <Synonymy:>/

#2. <Common name:>/

#3. Body length/

mm/

Data on body length is based largely on published descriptions, with the exception of several Anastrepha spp., Ceratitis rosa and Plioreocepta poeciloptera.

More significance should be attached to the maximum size of the larvae than to the minimum size, since it is possible that the latter represents third instar larvae which were not yet mature.

#4. Body <shape>/

1. slender, elongate, tapering anteriorly/

2. stout, elongate, tapering anteriorly/

3. elongate-cylindrical/

4. elongate, swollen anteriorly/

5. short-cylindrical or barrel-shaped/

6. pear-shaped/

States of this character represent points on a continuum. In general, among the fruit-infesting species, Bactrocera tend to be coded as state 1, Anastrepha as state 2 and Rhagoletis as state 3. Leaf-mining and stem-boring species often fit states 3 or 4, and species infesting Asteraceae (other than as miners) almost always fit states 5 or 6.

#5. Integument <sclerotization presence>/

1. unsclerotized, entirely whitish to yellowish/

2. partially sclerotized/

Sclerotization of the integument refers to parts of the integument not normally sclerotized in tephritid larvae. The cephalopharyngeal skeleton, anterior spiracles, and rimae of the posterior spiracular slits are normally sclerotized. If no other sclerotization exists, code the specimen as “entirely whitish to yellowish”.

Less obvious places to examine for sclerotization are the ventral prothorax and the area of the caudal segment between the intermediate tubercles (below the posterior spiracles).

Irregular sclerotization (of muscle scars in particular, or of other small, asymmetrically placed spots) in a few specimens of a sample, is abnormal, but not uncommon.

#6. Integument sclerotization <extent>/

1. forming a transverse line beneath posterior spiracles (mature larvae)/

2. covering entire caudal segment/

3. covering part of caudal segment/

4. covering part of dorsum/

5. covering part of dorsum and caudal segment/

6. forming a small heart-shaped patch of pigment below cuticle surface on ventral side of T1 (mature larvae)/

Mature larvae of several pest species, including Bactrocera cucurbitae, may have a transverse sclerotized line on the caudal segment between the tubercles below the posterior spiracles.

A heart-shaped pigmented area has been reported on the ventral side of the prothorax in B. umbrosa (White & Elson-Harris 1992).

Larvae with large sclerotized areas on the caudal segment and/or dorsum are not known to be pests of fruits; most attack Asteraceae, and one (Plioreocepta poeciloptera) bores in the stems of asparagus.

#7. Sclerotized process(es) on caudal segment <presence>/

1. absent/

2. present, strongly forked/

3. present, weakly forked/

4. present as two separate processes/

Sclerotized processes on the caudal segment are very rare in Tephritidae. They are present in Plioreocepta poeciloptera (strongly forked), Terellia (Cerajocera) ceratocera (weakly forked), and T. (C.) plagiata (two separate processes).

#8. Caudal ridge <presence>/

1. present/

2. absent/

The caudal ridge is diagnostic of Dacini and Ceratitini, although hard to see or possibly absent in Bactrocera (Tetradacus). It is a ridge of thickened cuticle in the intermediate region of the caudal segment below each posterior spiracle. It is usually readily distinguishable by SEM as well as under the dissecting microscope, but its presence or absence cannot generally be reliably determined in slide-mounted specimens (the presence of strong ridge-like structures in slide mounted specimens represents the collapse of a tubercle, not the caudal ridge of interest here; see dental sclerites instead).

Under the dissecting microscope, examine the caudal profile from all angles by rolling the specimen while it is completely submerged in alcohol, and you should be able to see the location of the ridge by the change in the light diffraction in the thickened part.

In species which have the intermediate tubercles I1 and I2 very pronounced (most Carpomyina; Euleia), there may also appear to be a ridge, but this is due to the proximity of the tubercles themselves, and not to a thickening of the cuticle between them (see also stomal guards).

#9. Mature larvae <jumping ability>/

1. unable to jump/

2. able to jump/

Some tephritids (Bactrocera, Ceratitis, Dacus, Euphranta, Blepharoneura) are known to “skip” or jump by grabbing their caudal segment with their mandibles and then suddenly releasing the tension, which propels them several inches away; larvae of most other genera (including Anastrepha, Toxotrypana, Rhagoletis and all tephritines) are not known to jump.

Head #10. Head <shape>/

1. of normal shape/

2. deflexed, laterally flattened, thorax hood-like/

Fruit-infesting species, as far as is known, have the head in anterior and frontal views rounded, and the dorsal anterior margin of the prothorax is not hood-like. Leaf-mining species, such as Euleia spp. and others, may have the head in anterior and/or frontal views laterally compressed, and the anterior margin of the thorax is extended anteriorly, hood-like, over the dorsal side of the head.

#11. Cephalic lobes <development>/

1. well developed/

2. moderately developed/

3. slightly developed/

4. not developed/

The cephalic lobes bear the antenna and the maxillary palp.

The figures illustrate the relative degree of development of the cephalic lobes typical of (1) Bactrocera and Dacus, (2) Anastrepha and Ceratitis, (3) Carpomya, Myiopardalis, and Rhagoletis, and (4) Euleia and Tephritinae.

Note that not all species are “typical” in the development of the cephalic lobes. Furthermore, species lacking well-developed cephalic lobes may differ substantially from the figure for state 4 in regards to the general appearance of the rest of the mask.

#12. Antenna <apparent segmentation>/

1. 1-segmented/

2. 2-segmented/

The antenna consists of a basal segment and an apical segment, both of which have sclerotized walls. Usually these two segments are clearly visible by SEM and on slide-mounted specimens. In species which bore or mine in stems and leaves, or infest flowerheads and galls, there appears to be only a single segment (by SEM) only slightly more pronounced than the surrounding reticulation; or the sclerotized part of the basal segment may be narrow and ringlike (in slide-mounted specimens), and not evident externally in these species.

#13. Stomal organ: primary lobe <size and shape>/

1. small, round/

2. large, elongate-rounded/

3. large, elongate-flattened/

4. rounded, protuberant/

The primary lobe of the stomal organ is the lobe which bears the stomal sensilla, located just anterior to the oral ridges, and ventral to the maxillary palp.

In Dacini, Ceratitini, and Tephritinae, it is small and round and surrounded by secondary lobes or reticulation. In Toxotrypanini, it is large and elongate-rounded. In leaf-mining and stem-boring species so far studied (e.g. Euleia), it is elongate-flattened, with the sensilla recessed in a pit. In the subtribe Carpomyina and in Euphrantinae it is rounded and protuberant.

#14. Stomal organ: number of peg sensilla <SEM>/

1. one/

2. two/

3. three <may be branched>/

4. four or more/

5. none apparent/

The stomal organ is located just anterior to the oral ridges, and ventral to the maxillary palp. The following sensilla are typically present on the stomal organ: 1 to 3 (or more) peg-like sensilla, 1 campaniform sensillum, and 2 pit sensilla. The peg sensilla are branched in some species, which may make them appear more numerous than they are. A single stout peg sensillum is characteristic of subtribe Carpomyina.

#15. Stomal organ: peg sensilla <branching>/

1. unbranched/

2. with few short branches/

3. with many long branches/

The peg sensilla are borne on the primary lobe of the stomal organ, which is located just anterior to the oral ridges, and ventral to the maxillary palp. They are sometimes branched, strongly so in Bactrocera tryoni, for example.

In B. dorsalis obtained from culture in Hawaii, the degree of branching was variable.

#16. Stomal organ: other peg-sensilla-like structures <SEM>/

1. present/

2. absent/

The stomal organ is located just anterior to the oral ridges, and ventral to the maxillary palp. The following sensilla are typically present in a slight depression on the stomal organ: 1 to 3 (or more) peg-like sensilla, 1 campaniform sensillum, and 2 pit sensilla. In three species of Anastrepha (A. leptozona, A. serpentina and A. striata), additional structures resembling peg sensilla surround this depression.

#17. Stomal region: secondary lobes <presence, shape>/

1. absent/

2. present, short, leaf-like/

3. present, medial ones elongate, like oral ridges/

4. embedded in overall reticulation/

The primary lobe of the stomal organ is the lobe which bears the stomal sensilla, and is located just anterior to the oral ridges, and ventral to the maxillary palp. The secondary lobes (when present) surround the small primary lobe, or may be reduced to a few antero-medial lobes. These can frequently be seen with a good dissecting microscope, using the same technique as for counting oral ridges, however SEM provides a much more detailed view.

Short leaf-like secondary lobes are typical of Ceratitini, and of the Bactrocera group of subgenera of Bactrocera. Long, ridge-like medial secondary lobes are typical of the Zeugodacus group of subgenera of Bactrocera, and are also present in Dirioxa; Dacus may also have this type of structure (see fig. of D. lounsburyii). Bactrocera oleae has round, protuberant secondary lobes.

Secondary lobes are absent in other genera of fruit flies, although the stomal organ of Tephritinae is often embedded in the overall reticulation of the head, some of which might be considered homologous to secondary lobes, This is also true of some other species which also inhabit the drier host microhabitats in flowerheads, leaves, stems and roots.

Secondary lobes which may subtend the primary lobe never bear sclerotized teeth (see stomal guards instead).

#18. Stomal region: number of secondary lobes <when present>/

Count all lobes which are closely associated with the primary lobe of the stomal organ.

#19. Stomal region: margins of secondary lobes <serration> <SEM>/

1. all entire/

2. one lobe with serrate margins/

3. more than one lobe with serrate margins/

Secondary lobes may surround the sensilla-bearing primary stomal lobe; their margins are usually entire, but in some species (e.g. Bactrocera latifrons) are distinctly dentate, serrate, or scalloped.

#20. Stomal region: sclerotized stomal guards <presence>/

1. present/

2. absent/

Sclerotized stomal guards are found as brown denticles on the posterior or ventral side of the primary lobe. They are present in many Carpomyini (Rhagoletis, Carpomya, Myiopardalis, Zonosemata), and appear as brown spot(s) or as a row of teeth when viewed with a dissecting microscope. In slide-mounted material they are frequently obscured by the mandibles. In SEM preparations, while the denticles per se are clearly visible, it is not known if they are all always sclerotized, but this seems to be the case. For purposes of this expert system, they have not been considered as “secondary lobes” of the stomal region.

#21. Stomal region: number of sclerotized stomal guards <when present>/

Sclerotized stomal guards are found as brown denticles on the posterior or ventral side of the primary lobe. They are present in many Carpomyini (Rhagoletis, Carpomya, Myiopardalis, Zonosemata), and appear as brown spot(s) or as a row of teeth when viewed with a dissecting microscope. In slide-mounted material they are frequently obscured by the mandibles. In SEM preparations, while the denticles per se are clearly visible, it is not known if they are all always sclerotized, but this seems to be the case. For purposes of this expert system, they have not been considered as “secondary lobes” of the stomal region.

#22. Oral ridges <presence>/

1. present/

2. absent/

Oral ridges are transverse ridges located on either side of the mandibles.

#23. Number of oral ridges <=ORL> <when present>/

Oral ridges are transverse ridges located on either side of the mandibles. They are best counted after first allowing the liquid to drain out of the grooves beneath them, if a dissecting microscope is used. Do not count the small detached accessory plates (q.v.) on the fringe of this area. (=ORL)

#24. Oral ridges margins <shape> <SEM>/

1. entire/

2. serrate <small teeth>/

3. dentate <large, sharply pointed teeth>/

4. scalloped <large, rounded teeth>/

5. emarginate <with rounded notches>/

6. occasionally incised/

7. rounded, more like reticulation than ridges/

Oral ridges are transverse ridges located on either side of the mandibles. Their margins are best seen under SEM. In slide-mounted material, margins usually appear entire.

#25. <Oral ridges> accessory plates <presence>/

1. present/

2. absent/

Accessory plates are found at the lateral margin of the oral ridges, and may be small and shell-like or elongate and imbricate with the oral ridges.

#26. <Oral ridges> number of accessory plates <when present>/

Accessory plates are found at the lateral margin of the oral ridges, and may be small and shell-like or elongate and imbricate with the oral ridges.

#27. <Oral ridges> accessory plates margins <whether serrated> <SEM>/

1. unserrated/

2. serrated/

Accessory plates are found at the lateral margin of the oral ridges, and may be small and shell-like or elongate and imbricate with the oral ridges. Their margins usually resemble those of the oral ridges.

#28. Elongate, finger-like lobes arising above mandibles <presence> <SEM>/

1. present/

2. absent/

Very elongate, finger-like lobes arising from the reticulation above the mandibles is common among Tephritinae, most of which infest Asteraceae. At present it is not known to occur among other subfamilies.

#29. Median oral lobe <presence, shape>/

1. protruding and laterally expanded/

2. protruding as a pair of flattened, 3- to 4-lobed structures/

3. absent or not protruding <as above>/

The median oral lobe lies in the preoral cavity between the mandibles. It is usually unremarkable, but in Bactrocera aquilonis it has been reported as protruding and laterally expanded (White & Elson-Harris 1992), and in Plioreocepta poeciloptera it is divided into a pair of flattened 3- to 4- lobed structures which lie along the medial face of the mandibles.

#30. Labium <width>/

1. broad/

2. narrow/

The labium is located on the ventral part of the head, posterior to the mandibles. As far as is known, it is broadly triangular in fruit-infesting species, but in leaf-mining or stem boring species, and in species infesting Asteraceae, it tends to be elongate and narrow.

Cephalopharyngeal skeleton #31. Mandibles: subapical teeth <presence>/

1. present/

2. absent/

Subapical teeth on the mandible are located on the ventral side of the mandible, sometimes on the lateral or medial edge, or both (when two teeth are present). Examine slide-mounted specimens carefully, focussing up and down, to determine the presence or absence of subapical tooth or teeth. These teeth are absent in most fruit-infesting species (use EXPA to see exceptions), and present in most leaf-mining or stem-boring trypetines and most, if not all, tephritines.

Note: previously published figures of Euleia fratria and Rhagoletis fausta (Phillips 1946) have omitted the well developed subapical teeth.

#32. Mandibles: subapical tooth <size>/

1. minute, visible only in slide mounts or by SEM/

2. much smaller than apical tooth, and delicate/

3. smaller than apical tooth, and very stout/

4. about the same size as apical tooth/

Subapical teeth are located on the ventral side of the mandible, sometimes on the lateral or medial edge, or both (when two teeth are present).

The size of the subapical tooth may vary, particularly where states 1 & 2 are concerned. It is not known if this is due to wear or represents intrinsic variation.

#33. Mandibles: <number and position of subapical tooth or teeth>/

1. with a single ventral tooth/

2. with a single medial tooth/

3. with a single lateral tooth/

4. with medial and lateral teeth/

Subapical teeth are located on the ventral side of the mandible, sometimes on the lateral or medial edge, or both (when two teeth are present).

#34. Mandibles: base <shape>/

1. stout, nearly perpendicular to a line from ventral part of base to apex of mandible/

2. elongate, forming a more oblique angle/

In Trypetinae and Tephritinae, the base of the mandibles tends to be stout, perpendicular, and does not form an elongate neck dorsally. In Dacinae, the dorsal part of the base is somewhat elongate, forming a neck, and the base of the mandibles is somewhat more oblique with respect to the tip of the mandibles.

#35. Parastomal bars <presence, shape>/

1. elongate, free from hypopharyngeal sclerite/

2. short and stout, or apparently absent/

The parastomal bars are generally readily visible in fruit-infesting Tephritidae as slender sclerites arising on the tentoropharyngeal sclerite and extending anteriorly dorsal to the hypopharyngeal sclerite. In leaf-mining and stem-boring trypetines, and in tephritines, they appear to be short, stout and so closely associated with the hypopharyngeal sclerite as to appear to be absent.

#36. Dental sclerites <presence, position>/

1. present, posterior to mandibles/

2. apparently absent, not visible in lateral view/

Dental sclerites are small, elongate sclerites immediately posterior or medial to the ventral part of the base of the mandibles. When they are posterior, as in Dacini, Ceratitini, and Dirioxa pornia, they are readily visible in slide preparations. If not readily visible they may be medial or absent, but they have not been studied sufficiently to distinguish between these two possibilities.

Spinules and creeping welts #37. Dorsal spinules on segments <distribution>/

1. T1/

2. T1-T2/

3. T1-T3/

4. T1-A1/

5. T1-A2/

6. T1-A3/

7. T1-A4/

8. T1-A5/

9. T1-A6/

10. T1-A7/

11. T1-A8/

12. with an unusual distribution/

The distribution of spinules on the body is best determined on slide-mounted material, however, other means can also be used. The number of rows of spinules on the anterior margin of the dorsum of each segment tends to become reduced in the more posterior segments, with the segments on which they last tend to appear being of some taxonomic use.

Caudal segment (a8) and anal lobes #38. Sensilla on caudal segment <A8> <number>/

1. 10 pairs, with at least 7 pairs visible under dissecting microscope <standard arrangement>/

2. 6 or fewer pairs visible/

The complete complement of sensilla on the caudal segment consists of: 2 dorsal (D1, D2); 1 lateral (L); 4 intermediate (I1a+b; I2; I3); and 3 ventral (V) for a total of 10 pairs of sensilla. No exceptions have been found among fruit-infesting species of Tephritidae. Some of these sensilla may be borne on minute papillae or tubercles, while others are visible (as in Toxotrypana) only with the greatest difficulty with the dissecting microscope. In slide-mounted material they often cannot be seen at all.

#39. Dorsal caudal sensilla <presence, arrangement>/

1. absent or visible only with great difficulty/

2. obvious, sessile or on separate papillae or tubercles/

3. obvious, on a single papilla or tubercle/

The two pair of dorsal caudal sensilla are located dorsal to the posterior spiracles.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means they are definitely identifiable as sensilla, although the novice is likely to consider that they hard to see.

“On separate papillae or tubercles” means they are visible as four separate small papilla scarcely larger than a sensillum, but easily seen.

“On a single papilla or tubercle” means they are visible as two bifid tubercles.

#40. Intermediate caudal sensillum I3 <presence, arrangement>/

1. absent or visible only with great difficulty/

2. obvious, but not on a papilla or tubercle/

3. obvious, on a papilla or tubercle/

The intermediate caudal sensillum I3 is located near, but slightly ventral to the lateral corner of the spiracular area.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means they are definitely identifiable as sensilla, although the novice is likely to consider that they hard to see.

“On a papilla or tubercle” means a papilla or very small tubercle scarcely larger than a sensillum, but easily seen.

#41. Lateral caudal sensillum <presence, arrangement>/

1. absent or visible only with great difficulty/

2. obvious, but not on a papilla or tubercle/

3. obvious, on a papilla or tubercle/

The lateral caudal sensillum L is located on the lateral part of the caudal segment about halfway between the spiracular area or the I3 sensillum, and the anterior margin of the segment.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means they are definitely identifiable as sensilla, although the novice is likely to consider that they hard to see.

“On a papilla or tubercle” means a papilla or very small tubercle scarcely larger than a sensillum, but easily seen.

#42. Intermediate caudal sensilla I1a&b <presence, arrangement>/

1. absent or visible only with great difficulty/

2. obvious, but not on a papilla or tubercle/

3. obvious, on a single papilla or tubercle, not bifid apically/

4. obvious, on a single papilla or tubercle, bifid apically/

The intermediate sensilla I1a and I1b are located very close together on the medial side of the intermediate area, below the posterior spiracles. When there is a caudal ridge (q.v.) these sensilla are at the medial corner of it.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means they (or at least one of them) are definitely identifiable as sensilla, although the novice is likely to consider that they hard to see.

“On a single papilla or tubercle, not bifid apically” means a small papilla scarcely larger than a sensillum, on which no more than one sensillum can be seen.

“On a single papilla or tubercle, bifid apically” means a small papilla scarcely larger than a sensillum, on which two sensilla can be seen.

#43. Intermediate caudal sensillum I2 <presence>/

1. absent or visible only with great difficulty/

2. obvious/

The intermediate sensillum I2 is located on the lateral side of the intermediate area. When there is a caudal ridge, the I2 sensillum is located below it.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means they are definitely identifiable as sensilla, although the novice is likely to consider that they hard to see; or they may be on a small papilla.

#44. Intermediate caudal sensilla I1a&b and I2 <arrangement>/

1. on separate papillae or tubercles/

2. on a single tubercle/

The intermediate sensilla I1a+b and I2, may be on separate and widely separated papillae (Anastrepha), or they may be somewhat closer together on the same tubercle (Rhagoletis, Carpomya).

#45. Ventral caudal sensilla <presence, arrangement>/

1. absent or visible only with great difficulty/

2. obvious, but not on a papilla or tubercle/

3. obvious, on a papilla or tubercle/

The three ventral caudal sensilla are located between the intermediate area and the anal area. Relative to the other sensilla they are usually more difficult to see, and some or all of them may be reduced to mere pits. However, in the Carpomyina (Rhagoletis, Carpomya etc.), they are on a large tubercle similar to the intermediate tubercle.

“Absent or visible only with great difficulty” means the sensilla are practically indistinguishable from the specks of dust that are inevitably floating in the preservation fluid.

“Obvious” means at least one is identifiable as a sensillum, although the novice is likely to consider that it is hard to see.

“On a papilla” means a small papilla scarcely larger than a sensillum, but easily seen.

“On a tubercle” means a low protuberance of much larger circumference than a papilla.

#46. Anal lobes <prominence>/

1. very protuberant/

2. plainly visible, but not strongly protuberant/

3. indistinct, retracted into anal slit, or absent/

The anal lobes usually are plainly visible in Tephritidae, particularly among fruit-infesting species. Among these a few species (e.g. Rhagoletis cerasi) have the anal lobes very protuberant and somewhat expanded distally. The degree of protrusion, however, may vary among individuals: in life, they are capable of being protruded and retracted somewhat. In most Tephritinae and certain other species the anal lobes may be absent, indistinct, or retracted within the anal slit.

#47. Anal lobes <shape>/

1. simple/

2. grooved/

3. bifid/

The anal organ consists of a pair of lobes or perianal pads. Each of these lobes may be entire, grooved, or distinctly bilobed.

Anterior spiracles #48. Anterior spiracle <shape in profile>/

1. elevated, margin convex to straight/

2. elevated, margin concave medially, appearing bilobed/

3. not strongly elevated, margin long and convex/

This character refers to the outline of the external part of the anterior spiracle, across the tubules. In states 1 and 2, the tubules are somewhat elevated above the level of the surrounding cuticle. In state 3, the tubules are within a sulcus or nearly covered by the anterior margin of the mesothoracic segment. In slide mounted specimens, state 3 is represented by having the internal basal part of the spiracle often not preserved, leaving only the broad arc of tubules on the distal part.

#49. Anterior spiracular tubules <number> <=ANS>/

Anterior spiracular tubules (digits) are best counted after first allowing the liquid to evaporate somewhat from the body of the larva, if a dissecting microscope is used. A compound microscope is sometimes better, but requires slide preparation. (=ANS)

#50. Anterior spiracular tubules <arrangement>/

1. in a single uniform row/

2. in a single irregular row/

3. in two rows/

4. in three or more rows/

Anterior spiracular tubules are most often in a single uniform or irregular row. Two rows occur in some Rhagoletis, and Plioreocepta poeciloptera has more than three rows.

Posterior spiracles #51. Posterior spiracular area <whether set off>/

1. set off by a ring of tubercles and a distinct groove/

2. not distinctly set off from caudal segment/

Euleia spp. and at least some other Trypetini have the posterior spiracles surrounded by extremely well developed dorsal and intermediate tubercles which form a ring around the spiracular area that distincly sets this area off from the rest of the caudal segment.

#52. Posterior spiracles: average length of dorsal and ventral rimae <=LTH>/

µm/

Measure the rimae from the outside edges. For Anastrepha species which are difficult to distinguish, take the average of the dorsal and ventral rimae. (=LTH)

#53. Posterior spiracles: slits <length:width ratio>/

x longer than wide/

Measure the length and width of the rimae from the outside edges and take the ratio of length to width. For species other than Anastrepha, much of the data in the system taken from the literature appears to have been “eyeballed”.

#54. Dorsal spiracular processes <number and arrangement of trunks>/

1. with numerous trunks arising from an elongate base/

2. with numerous trunks arising from a short or semicircular base/

3. with a few trunks radiating from a short or semicircular base/

4. with only 1 or 2 trunks/

The form of the spiracular processes is a useful character for comparative purposes, but is difficult to encode as a single useful character for matching taxa in the database. The four states presently given for the dorsal (SPI) spiracular processes represent only two aspects of their form.

#55. Number of dorsal spiracular processes/

Count the number of trunks of spiracular processes in the dorsal group. The dorsal group can usually be identified even in disarticulated slide mounts of the caudal segment: it is the group oriented approximately at right angles (rather than at approximately the same angle) to the lateral group next to it.

#56. Number of ventral spiracular processes/

Count the number of trunks of spiracular processes in the ventral group. The ventral group can usually be identified even in disarticulated slide mounts of the caudal segment: it is the group oriented approximately at the same angle (rather than at approximately a right angle) as the lateral group next to it.

#57. Number of lateral spiracular processes <on each side>/

Count the number of trunks of spiracular processes in each lateral group. You can enter the data as a range of values, or as the average for the two groups. (Data in the system is entered as a range of values). For example, if group 2 has 6 trunks and group 3 has 10 trunks, enter the data as 6–10 or as 8. These two methods may retrieve slightly different groups of taxa due to the way the program handles numeric data entered as ranges.

#58. Basal width of spiracular processes <=BAS>/

µm/

Measure the basal width of the spiracular processes as the distance between outer-most trunks at their insertion point. This ratio is useful in discriminating among certain species of Anastrepha. (=BAS)

#59. Dorsal and ventral spiracular processes: average number of tips <=TIP>/

Count the number of tips of the spiracular processes in the dorsal and ventral groups and take the average.

#60. Dorsal and ventral spiracular processes: ratio of number of tips to number of trunks <=RTO2>/

The branching ratio is the ratio of the number of tips to number of trunks. This ratio is useful in discriminating among certain species of Anastrepha. (=RT02)

#61. Area between posterior spiracles <texture>/

1. smooth/

2. with a cellular or rugose appearance/

Carpomya vesuviana, C. schineri (not currently in the system) and Myiopardalis pardalina have the area between the posterior spiracles with a rugose, cellular or alveolate appearance. Allow the surface of the caudal segment to dry briefly to better observe this character.

Linear descriminant functions #62. Anastrepha striata vs. A. bistrigata: C = 23.5log(BAS) – 0.75(ANS) – 0.63(TRK) – 15:/

1. C greater than 0 <A. striata>/

2. C less than 0 <A. bistrigata>/

LINEAR DISCRIMINANT FUNCTION

To distinguish between Anastrepha striata and A. bistrigata, insert the values for BAS, ANS and TRK, and calculate C.

C = 23.5log(BAS) – 0.75(ANS) – 0.63(TRK) – 15

where
ANS = the number of tubules on the anterior spiracle,
BAS = the basal width of the spiracular processes as the distance between outer-most trunks at their insertion point,
TRK = the number of trunks on the posterior spiracular processes.

#63. Anastrepha fraterculus vs. A. obliqua: C = 22.70log(BAS) – 0.45(LTH) – 0.99(ANS) + 24.00:/

1. C greater than 0 <A. fraterculus>/

2. C less than 0 <A. obliqua>/

LINEAR DISCRIMINANT FUNCTION

To distinguish between Anastrepha fraterculus and A. obliqua, insert the values for BAS, LTH and ANS, and calculate C.

C = 22.70log(BAS) – 0.45(LTH) – 0.99(ANS) + 24.00

where
ANS = the number of tubules on the anterior spiracle,
BAS = the basal width of the spiracular processes as the distance between outer-most trunks at their insertion point,
LTH = the length of the rimae of the posterior spiracular openings.

#64. Anastrepha obliqua vs. A. suspensa: C = 1.40(ANS) + 0.60(TIP) + 0.24(LTH) – 2.45(ORL) – 30.93:/

1. C greater than 0 <A. obliqua>/

2. C less than 0 <A. suspensa>/

LINEAR DISCRIMINANT FUNCTION

To distinguish between Anastrepha obliqua and A. suspensa, insert the values for ANS, TIP, LTH and ORL, and calculate C.

C = 1.40(ANS) + 0.60(TIP) + 0.24(LTH) – 2.45(ORL) – 30.93

where
ANS = the number of tubules on the anterior spiracle.
LTH = the length of the rimae of the posterior spiracular openings.
ORL = the number of oral ridges (per side).
TIP = the number of branch tips of the dorsal or ventral posterior spiracular processes.

#65. Anastrepha fraterculus vs. A. suspensa: C = 24.20log(BAS) + 26.67log(RTO2) – 40.52:/

1. C greater than 0 <A. fraterculus>/

2. C less than 0 <A. suspensa>/

LINEAR DISCRIMINANT FUNCTION

To distinguish between Anastrepha fraterculus and A. suspensa, insert the values for BAS and RTO2, and calculate C.

C = 24.20log(BAS) + 26.67log(RTO2) – 40.52

where
BAS = the basal width of the spiracular processes as the distance between outer-most trunks at their insertion point.
RTO2 = the ratio of the number of branch tips to the number of trunks of the posterior spiracular processes.

#66. Anastrepha suspensa vs. A. fraterculus vs. A. obliqua: Cf = 186.07log(BAS) + 13.01(ORL) + 39.58log(RTO2) + 8.63(ANS) + 1.10(LTH) – 294.42; Cs = 151.06log(BAS) + 17.23(ORL) – 5.62log(RTO2) + 8.51(ANS) + 1.25(LTH) – 285.19; Co = 161.23log(BAS) + 14.17(ORL) + 27.96log(RTO2) + 9.77(ANS) + 1.58(LTH) – 326.32:/

1. Cf is largest value <A. fraterculus>/

2. Cs is largest value <A. suspensa>/

3. Co is largest value <A. obliqua>/

LINEAR DISCRIMINANT FUNCTION

To distinguish between Anastrepha suspensa, A. fraterculus and A. obliqua, insert the values for BAS, ORL, RTO2, ANS and LTH, and calculate Cf, Cs, and Co.

Cf = 186.07log(BAS) + 13.01(ORL) + 39.58log(RTO2) + 8.63(ANS) + 1.10(LTH) – 294.42

Cs = 151.06log(BAS) + 17.23(ORL) – 5.62log(RTO2) + 8.51(ANS) + 1.25(LTH) – 285.19

Co = 161.23log(BAS) + 14.17(ORL) + 27.96log(RTO2) + 9.77(ANS) + 1.58(LTH) – 326.32

where
ANS = the number of tubules on the anterior spiracle.
BAS = the basal width of the spiracular processes as the distance between outer-most trunks at their insertion point.
LTH = the length of the rimae of the posterior spiracular openings.
ORL = the number of oral ridges (per side).
RTO2 = the ratio of the number of branch tips to the number of trunks of the posterior spiracular processes.

#67. <Host plants:>/

1. Actinidiaceae/

2. Agavaceae/

3. Anacardiaceae/

4. Annonaceae/

5. Apiaceae <=Umbelliferae>/

6. Apocynaceae/

7. Aquifoliaceae/

8. Araliaceae/

9. Araucariaceae/

10. Arecaceae/

11. Asclepiadaceae/

12. Asteraceae <=Compositae>/

13. Berberidaceae/

14. Bignoniaceae/

15. Bombacaceae/

16. Brassicaceae <=Cruciferae>/

17. Bromeliaceae/

18. Cactaceae/

19. Canellaceae/

20. Capparaceae <=Capparidaceae>/

21. Caprifoliaceae/

22. Caricaceae/

23. Caryophyllaceae/

24. Cecropiaceae/

25. Celastraceae <=Siphonodontaceae>/

26. Chrysobalanaceae/

27. Clusiaceae/

28. Combretaceae/

29. Convolvulaceae/

30. Cornaceae/

31. Cucurbitaceae/

32. Cunoniaceae/

33. Cupressaceae/

34. Davidsoniaceae/

35. Ebenaceae/

36. Elaeocarpaceae/

37. Eleagnaceae/

38. Ericaceae/

39. Erythroxylaceae/

40. Euphorbiaceae/

41. Fabaceae <=Leguminosae>/

42. Fagaceae/

43. Flacourtiaceae/

44. Gnetaceae/

45. Grossulariaceae/

46. Juglandaceae/

47. Lamiaceae <=Labiatae>/

48. Lauraceae/

49. Lecythidaceae <=Barringtoniaceae>/

50. Liliaceae/

51. Loganiaceae/

52. Loranthaceae/

53. Malpighiaceae/

54. Malvaceae/

55. Meliaceae/

56. Moraceae/

57. Musaceae/

58. Myristicaceae/

59. Myrtaceae/

60. Olacaceae/

61. Oleaceae/

62. Opiliaceae/

63. Orchidaceae/

64. Orobanchaceae/

65. Oxalidaceae/

66. Paeoniaceae/

67. Passifloraceae/

68. Poaceae/

69. Podocarpaceae/

70. Polygonaceae/

71. Proteaceae/

72. Punicaceae/

73. Rhamnaceae/

74. Rhizophoraceae/

75. Rosaceae/

76. Rubiaceae <=Naucleaceae>/

77. Rutaceae/

78. Salicaceae/

79. Santalaceae/

80. Sapindaceae/

81. Sapotaceae/

82. Simaroubaceae/

83. Smilacaceae/

84. Solanaceae/

85. Staphyleaceae/

86. Sterculiaceae/

87. Symplocaceae/

88. Theaceae/

89. Tiliaceae/

90. Ulmaceae/

91. Urticaceae/

92. Verbenaceae/

93. Vitaceae/

94. Zygophyllaceae/

Host plants #68. <Host plant family (A) (see Notes):>/

1. Actinidiaceae/

2. Agavaceae/

3. Anacardiaceae/

4. Annonaceae/

5. Apiaceae <=Umbelliferae>/

6. Apocynaceae/

7. Aquifoliaceae/

8. Araliaceae/

9. Araucariaceae/

10. Arecaceae/

11. Asclepiadaceae/

12. Asteraceae <=Compositae>/

13. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

#69. <Host plant family (B-Ca) (see Notes):>/

1. Berberidaceae/

2. Bignoniaceae/

3. Bombacaceae/

4. Brassicaceae <=Cruciferae>/

5. Bromeliaceae/

6. Cactaceae/

7. Canellaceae/

8. Capparaceae <=Capparidaceae>/

9. Caprifoliaceae/

10. Caricaceae/

11. Caryophyllaceae/

12. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

Barringtoniaceae: see Lecythidaceae.

#70. <Host plant family (Ce-Cz) (see Notes):>/

1. Cecropiaceae/

2. Celastraceae <=Siphonodontaceae>/

3. Chrysobalanaceae/

4. Clusiaceae/

5. Combretaceae/

6. Convolvulaceae/

7. Cornaceae/

8. Cucurbitaceae/

9. Cunoniaceae/

10. Cupressaceae/

11. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

Compositae: see Asteraceae.

Cruciferae: see Brassicaceae.

#71. <Host plant family (D-K) (see Notes):>/

1. Davidsoniaceae/

2. Ebenaceae/

3. Elaeocarpaceae/

4. Eleagnaceae/

5. Ericaceae/

6. Erythroxylaceae/

7. Euphorbiaceae/

8. Fabaceae <=Leguminosae>/

9. Fagaceae/

10. Flacourtiaceae/

11. Gnetaceae/

12. Grossulariaceae/

13. Juglandaceae/

14. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

#72. <Host plant family (L-M) (see Notes):>/

1. Lamiaceae <=Labiatae>/

2. Lauraceae/

3. Lecythidaceae <=Barringtoniaceae>/

4. Liliaceae/

5. Loganiaceae/

6. Loranthaceae/

7. Malpighiaceae/

8. Malvaceae/

9. Meliaceae/

10. Moraceae/

11. Musaceae/

12. Myristicaceae/

13. Myrtaceae/

14. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

Leguminoseae: see Fabaceae.

#73. <Host plant family (N-P) (see Notes):>/

1. Olacaceae/

2. Oleaceae/

3. Opiliaceae/

4. Orchidaceae/

5. Orobanchaceae/

6. Oxalidaceae/

7. Paeoniaceae/

8. Passifloraceae/

9. Poaceae/

10. Podocarpaceae/

11. Polygonaceae/

12. Proteaceae/

13. Punicaceae/

14. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

Naucleaceae: see Rubiaceae.

#74. <Host plant family (R-Sa) (see Notes):>/

1. Rhamnaceae/

2. Rhizophoraceae/

3. Rosaceae/

4. Rubiaceae <=Naucleaceae>/

5. Rutaceae/

6. Salicaceae/

7. Santalaceae/

8. Sapindaceae/

9. Sapotaceae/

10. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

#75. <Host plant family (Sb-Z) (see Notes):>/

1. Simaroubaceae/

2. Smilacaceae/

3. Solanaceae/

4. Staphyleaceae/

5. Sterculiaceae/

6. Symplocaceae/

7. Theaceae/

8. Tiliaceae/

9. Ulmaceae/

10. Urticaceae/

11. Verbenaceae/

12. Vitaceae/

13. Zygophyllaceae/

14. none of the above/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

Siphonodontaceae: see Celastraceae.

Umbelliferae: see Apiaceae.

#76. <Notes on host plant:>/

Don't use more than 1 of the “host plant family (...)” characters in an identification.

#77. Part of plant attacked:/

1. fruit/

2. leaf/

3. stem/

4. root crown or stem base/

5. bud or flower other than capitulum/

6. capitulum (of Asteraceae)/

7. receptacle (see also ‘capitulum’)/

8. multiple seeds (including achenes; see also ‘capitulum’)/

9. single seed (including achene; see also ‘capitulum’)/

Biogeographic region #78. <Biogeographic region:>/

1. Nearctic/

2. Neotropical/

3. Palearctic/

4. Afrotropical/

5. Oriental/

6. Australasian-Oceanian/

For specimens from near regional boundaries, select as many regions as might apply.

Specimens examined #79. <Specimens examined or source of description:>/

#80. Sources of data and SEM numbers:/

Miscellaneous #81. Abbreviated name:/

#82. <Illustrations:>/


Cite this publication as: ‘L.E. Carroll, A.L. Norrbom, M.J. Dallwitz, and F.C. Thompson. 2004 onwards. Pest fruit flies of the world – larvae. Version: 8th December 2006. http://delta-intkey.com’.

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