The genus Carpomya Costa (Diptera: Tephritidae): new synonymy, description of first American species, and phylogenetic analysis

10 Pages·1997·4.4 MB·

by  A L Norrbom

PROC. ENTOMOL. SOC. WASH.
99(2), 1997, pp. 338-347
THE GENUS CARPOMYA COSTA (DIPTERA: TEPHRITIDAE):
NEW SYNONYMY, DESCRIPTION OF FIRST AMERICAN SPECIES,
AND PHYLOGENETIC ANALYSIS
Allen L. Norrbom
Systematic Entomology Laboratory, PSL Agricultural Research Service, U.S. Depart-
ment of Agriculture % National Museum of Natural History, MRC 168, Washington, DC
20560, U.S.A.
—
Abstract. Carpomya tica, new species, from Costa Rica is described and reported as
the first species of Carpomya from the Americas. Phylogenetic relationships within Car-
pomya are analyzed. Myiopardalis Bezzi and Goniglossum Rondani are recognized as
subjective junior synonyms of Carpomya. Carpomya wiedemanni, n. comb., and C. par-
dalina (originally described in Carpomya) are transferred to Carpomya from Goniglossum
and Myiopardalis, respectively, and Spilographa caucasica Bigot is removed from the
genus. A lectotype is designated for C pardalina, and a key to the species of Carpomya
is provided.
—
Resumen. Se describe Carpomya tica, especie nueva, de Costa Rica, la primera es-
pecie de Carpomya de las Americas. Se analizan las relaciones filogeneticas dentro de
Carpomya. Myiopardalis Bezzi y Goniglossum Rondani son sinonimos subjectivos nuevos
de Carpomya. Se transfieren C. pardalina (descrita originalmente en Carpomya) y Car-
pomya wiedemanni, n. comb., a Carpomya de Myiopardalis y Goniglossum, respectiva-
mente, y se remueve Spilographa caucasica Bigot de este genero. Se designa un lectotipo
para C. pardalina, y se provee una clave a las especies de Carpomya.
Key Words: Fruit flies, Tephritidae, Carpomya, Myiopardalis, Goniglossum
The genus Carpomya Costa (1854) pre- Methods
viously included four species known from j f^n^^ j^e morphological terminology
the southern Palearctic, northeastern Afro- ^f McAlpine (1981), except as noted by
tropical, and western Oriental Regions, al- Norrbom and Kim (1988). Terminology for
though one of these species, C. caucasica (he wing pattern follows Foote (1981, see
(Bigot) does not seem to belong in this ge- pjg 71) Taxonomically useful characters
nus. The monotypic genera Myiopardalis vvithin Carpomya are listed in Table 1 and
Bezzi (1910) and Goniglossum Rondani the distributions oftheir states are shown in
(1856), here considered new subjective ju- Table 2, but only the nine characters with
nior synonyms of Carpomya, included one an asterisk were used in the phylogenetic
southern Palearctic/westem Oriental species analysis; the others are autapomorphies.
and one Palearctic species, respectively. The Hennig86 program was used for phy-
This paper reports the discovery of the first logenetic analysis, with the Rhagoletis ta-
American species of Carpomya, increasing bellaria group as the outgroup for assigning
the number of species of this genus to six, character polarities. Transformation series
and substantially extending its distribution, and the polarity of some characters are fur-
.
VOLUME 99, NUMBER 2 339
Table 1. Characters taxonomically useful for species of Carpomya. Only characters with an * were used in
phylogenetic analysis.
—
1 Head elongate, gena especially produced and angulate anteriorly, and proboscis long geniculate 0) no;
1) yes. —
2. Facial carina 0) moderately to strongly produceddorsally, gradually becomingbroaderand lessproduced
ventrally; 1) weakly produced, gradually becoming broader ventrally; 2) strongly produceddorsally, rather
abruptlybecoming verybroadand weaklyproducednearmidlength; 3)extremelyproducedandmoderately
broad throughout, carinate on both sides. States 1, 2, and 3 are here considered independently derived
autapomorphies.
—
*3. Ocellar setae 0)—well developed; 1) minute, length less than width ofocellar tubercle.
4. First flagellomere 0) with dorsoapical p—oint; 1) rounded dorsoapically.
*5. Mesonotal color and microtrichia pattern 0) unicolorous except white areas, or ifpartially brown without
microtrichia or with microtrichia unicolorous; 1) with pattern of dark brown or black marks (similar to
Fig. 1), some without—microtrichia, others with dense black microtrichia.
*6. Scutellar color pattern 0) brown basally, white apically; 1) with medial brown spot extended to apex or
with smaller isolated medial brown spot and apical brown spot(s); 2) without medial brown spot, palearea
m-shaped, with medial pale arm; 3) entirely yellow. The transformation series ofthischaracterisuncertain
and it was coded unordered in the Hennig86 analysis. Carpomya tica, pardalina and wiedemanni were all
coded 1, although there are some differences in their scutellar patterns: C tica has a large medial brown
C
spot and U-shaped yellow area which includes the apical setae (Fig. 1); pardalina has a smallermedial
brown spot, isolated by a larger pale area from an apical brown spot or spots on which the apical setae
are located; and C. wiedemanni is variable, either like C pardalina, or with the medial brown spot large
and including th—e apical setae (Freidberg & Kugler 1989: 191).
*7. Scutellumshape 0) relatively triangularin dorsal view, disc flatto slightlyconvex; 1) almostsemicircular
in dorsal view, disc slightly to moderately convex (a little less convex in incompleta than in schineri and
vesuviana); 2)—almost semicircular in dorsal view, disc very strongly convex.
*8. Katepistemum 0) unicolorous brown {R. tabellaria group) or yellow (C. tica); 1) yellow ororange, with
dorsal margin white (contrast often weak in C. incompleta andpardalina); 2) with large brown or black
area, dorsal margin white. State appears to be plesiomorphic, although it should be noted that states 1
and 2 occur in Zonose—mata.
*9. Cell r, with spur vein 0) with at most a dark spot or crease within the subapical band; 1) with a distinct
spur vein origi—nating from vein R,+3 in the subapical band.
10. Crossvein r-m 0) near midpoint between bm-cu and dm-cu, within or at apical margin ofdiscal band; 1)
at 0.73 distance from bm-c—u to dm-cu, touching proximal margin ofsubapical band.
11. Subbasal and discal bands—0) not connected or connected posteriorly; 1) connected in cells r, and br.
12. Anterior ap—ical wing band 0) present; 1) absent.
*13. Epandrium 0)mostlybrown; 1)mostlyyellowwithsmalldorsomedialbrownspot. Inothermostlyyellow
Carpomyini the epandrium is mostly brown or all yellow or has paired dorsolateral spots, although in
Zonosemata—it varies from states to 1, sometimes intraspecifically.
14. Aculeus tip 0) nonse—rrate; 1) serrate.
*15. Spermatheca number 0) 3; 1)2. The number ofspermathecae varies from 2-3 in the R. tabellaria group
and other Rhagoletis and Carpomyini (Bush 1966, Norrbom 1994b), but 3 is the plesiomorphic state for
Tephritidae, so I tentatively coded that number as plesiomorphic within Carpomya and assigned that state
to the outgroup in —the Hennig86 analysis.
16. Spermathec—a shape 0) single chambered; 1) double chambered.
*17. Host plant 0) Rosa spp. (Rosaceae); 1) Ziziphus spp. (Rhamnaceae); 2) Cucurbitaceae (C paradalina
breeds in Cucumis, Citrullus. and Ecballium spp., and C wiedemanni inBryonia spp.). The transformation
series is uncertain; states 1 and 2 are probably apomorphic, as C. pardalina and C. wiedemanni are the
only two species of Carpomyini known to breed in Cucurbitaceae, and C. incompleta and C vesuviana
aretheonlyCarpomyini known from Rhamnaceae (Freidberg& Kugler 1989,White&Elson-Harris 1992),
but these states may be independently derived from state or some other state (a wide range of host
families are attacked by Carpomyini). The hosts ofC tica are unknown. Codedunordered in theHennig86
analysis.
340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Character state distributions among species of Carpomya.
——
VOLUME NUMBER
99, 2 341
Pakistan). In the Bigot Collection, now at
the University Museum, Oxford, there are
three specimens (2 6 1 9) on separate pins,
one of each sex also with a larva pinned
below the adult specimen. The pin of the
female, here designated as lectotype, has a
thin paper label of the style that Ackland
and Taylor (1972, Fig. 6) found mainly on
specimens with Bigot manuscript names.
The lectotype's label has the following data
in Bigot's writing: "Carpomyia pardalina
[the second half of the specific name writ-
ten over some other letters] 9 \n. sp. Inedit.
\ qui[rest of word illegible] Octobre 1890.
J. Bigot \ Beloutchistan \ attaque les mel-
ons." There is a fourth pin with a similar
label except for a "cJ" symbol instead of
"9." It has only some debris, perhaps the
remains of a third syntype male. The lec-
totype is pinned through its abdomen and
has slightly shriveled eyes, but otherwise is Fig. 1. Carpomya tica. thorax, dorsal view.
in good condition. All of these specimens
fit Bigot's description and the traditional in-
terpretation of this species. date of description, but their identity helps
Carpomyia caucasica Zaitzev (1919) to confirm the synonymy of this name.
was described from male and female spec- As explained in the Relationships sec-
imens from "East Transcausia, Dzhevan- tion, I follow Zaitzev (1919, 1947), Roh-
shir, Areshsk territory, Elisavetn region and dendorf (1939) and Kandybina (1965) in
North Mugan." Elisavetn is probably the including pardalina in Carpomya, rather
locality in Azerbaijan that has also been than in the monotypic genus Myiopardalis,
known as Gandzhe, Kirovabad, and Yeli- as it has been treated by the majority of
savetpol (A. Konstantinov, personal com- authors.
munication). This name has long been con-
sidered a synonym of C. pardalina by Rus- Carpomya tica Norrbom, new species
sian workers (Stackelberg 1928, Zaitzev (Figs. 1-4)
1947, Kandybina 1965), but has been little Holotype. S (at USNM, for eventual
noticed by western authors. It was omitted deposit in Instituto Nacional de Biodiver-
from the Palearctic Diptera catalog (Foote sidad, Heredia, Costa Rica), COSTA RICA:
1984). I examined a pair of specimens of San Jose: Zurqui de Moravia, 10°03'N
C. pardalina in the Zoological Institute, St. 84°0rw, 1600 m. Malaise trap, V.1992, P
Petersburg with the following label data: Hanson.
"A3, o. 3. CT." [= Azerbaijan?] and hand- Diagnosis. Carpomya tica differs from
written localities that I cannot decipher; the all other species of Carpomya and most
male with the date "19/viii 1928", and the other Carpomyina in the distal location of
female with the date "3/viii 1927." The fe- crossvein r-m (Fig. 2), which is at the apical
male also has a determination label with % of cell dm versus near the middle of that
''Myiopardalis caucasica m., Zaitzev det." cell in other Carpomyina except Crypto-
These specimens cannot be types because dacus spp. and Haywardina cuculi (Hen-
their collection dates are subsequent to the del), which differ from Carpomya in having
342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Carpomya tica, right wing.
C
a white medial scutal stripe or spot. tica notal lobe, dorsal margin of anepistemum,
also differs from other species of Carpo- and posterior third of notopleuron white.
myina, including most of the other species Mesonotum (Fig. 1) with following dark
of Carpomya, by the strongly swollen brown markings: medial spot from level of
shape ofits scutellum. Only C. schineri and dorsocentral seta to level ofacrostichal seta,
C. vesuviana approach C. tica in scutellum extended laterally slightly beyond acrosti-
shape, but neither has this structure as stout chal seta; spot on anterior two-thirds ofno-
as in C. tica, and they further differ in the topleuron, extended mesally on scutum to
shape of the pale area of their scutellar col- level of presutural supra-alar seta but not
or patterns, which is somewhat m-shaped, including it; broad spot along transverse su-
with a medial pale mark (see Freidberg and ture; spot from wing base to slightly mesal
Kugler 1989, Fig. 188), rather than U-shaped to level of intra-alar seta and from postsu-
as in C. tica (Fig. 1; the yellow areas are tural supra-alar seta almost to post-alar and
continuous on the extreme apex of the scu- intra-alar setae; spot posterior and lateral to
tellum, visible in posterior view). In the key intra-alar seta; and scutellum, except for
to Neotropical genera of Tephritidae in narrow, U-shaped yellow mark that in-
Foote (1980), C. tica will key to Cecido- cludes apical and basal setae. Pleuron with-
chares, which is not closely related (it be- out dark brown markings. Subscutellum en-
longs to the subfamily Tephritinae). Species tirely dark brown. Mediotergite yellow lat-
of the latter genus differ as follows: body erally, broadly dark brown medially, more
mostly dark brown, including all ofthe scu- broadly so dorsally. Scutellum strongly
tum and scutellum; at least some scutal swollen. Chaetotaxy as in Rhagoletis and
setulae swollen; and outer andinner surstyli Carpomya. Dorsocentral seta slightly ante-
short. — rior to level of postsutural supra-alar seta.
Description. Setae black. Body pre- Scapular setae white. Mesonotal setulae
dominantly yellow. Head: Yellow. 3 fron- slender; those on scutum yellow or black,
tal, 2 orbital setae, all large. Ocellar seta occurring in patches. Scutellum with yellow
large (length about equal to width offrons). setulae, present only on yellow areas.
Facial carina weak. First flagellomere about Mesonotum mostly moderately densely
2 times as long as wide (lateral view), with pale microtrichose, except for dark brown
small dorsoapical point. Proboscis short, part of notopleuron, dark brown spot along
capitate. Thorax: Mostly yellow. Postpro- transverse suture, lateral halfofbrown spot
I
VOLUME NUMBER
99, 2 343
mm
0.1
Figs. 3-4. Carpomya tica, male genitalia. 3, Epandrium and surstyli, posterior view (cerci not shown). 4,
Epandrium, surstyli and cercus, lateral view.
between postsutural supra-alar and intra- band in cell r,, and separated from costa by
alar setae, and spot posterior and lateral to hyaline marginal spots in cells rj and cell
intra-alar seta, which are densely black mi- r2+3. Cell r, without spur vein from vein
crotrichose, and the following nonmicrotri- R2+3 in subapical band. Crossvein r-m at
chose areas: dark brown part of scutum 0.73 distance from bm-cu to dm-cu (just
mesal to notopleural dark area; middle of touching proximal margin of subapical
medial scutal brown spot; mesal half of band). Cell bcu with apical lobe 0.5 times
brown spot between postsutural supra-alar as long as maximal width ofcell.Abdomen:
and intra-alar setae; and scutellum except Mostly yellow, each tergite (through tergite
yellow areas and base and apex of lateral 5) with unpaired medial and paired subla-
brown area. Subscutellum and upper halfof teral brown spots, small on syntergite 1+2.
brown area of mediotergite densely black- Male terminalia (Figs. 3-4): Epandrium
ish microtrichose; yellow parts of medi- brown, surstyli yellow. Outer surstylus
otergite moderately pale microtrichose; slender, with distinct mesal lobe; part apical
ventral part of brown area bare. Legs: Yel- to mesal lobe with relatively few small
low, except hind tibia with basal posterior setulae.
and subapical anterior and posterior brown
spots. Wing (Fig. 2): With subbasal, discal, Spilographa caucasica Bigot
preapical and anterior apical bands. Acces- Hendel (1927) placed this species in Car-
sory costal band and posterior apical bands pomya, but probably based on confusion
absent. Discal and subapical bands parallel, with Carpomyia caucasica Zaitzev (= C.
perpendicular to long axis of wing; each pardalina). Without examination of the ho-
broad, but not connected. Anterior apical lotype it is difficult to recognize S. caucas-
band only narrowly connected to subapical ica. Bigot's description is vague and there
344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
are no illustrations. But it is very doubtful pomyina, but other character state distri-
that S. caucasica belongs in Carpomya, as butions support that hypothesis (Norrbom
no species of that genus has a longitudinal 1989, 1994b).
band in addition to four transverse bands on Carpomya, as recognized here, includes
the wing. Bigot's statement "quatre bandes six species: C. incompleta (Becker), C. par-
roussatres, peu distinctes, sises a la partie dalina Bigot, C. schineri (Loew), C. vesu-
posterieure du thorax" (4 reddish bands, viana Costa, C wiedemanni (Meigen), New
C
not very distinct, situated at the posterior Combination and tica, n. sp. Carpomya
part of the thorax), also seems questionable tica belongs in the Carpomyina, or at least
for a species of Carpomya, which except to the Carpomyini, based on the shape of
C
for incompleta (which has only 3 wing its male genitalia; its surstyli have the typ-
bands and therefore cannot be S. caucasi- ical carpomyine shape (Figs. 3-4). With
ca), have distinct dark brown spots or larger most of the other species of Carpomya, C.
marks on the mesonotum. I cannot recog- tica shares a distinctive, apomorphic color
nize S. caucasica, and treat it here as an pattern ofthe cuticle and microtrichia ofthe
unplaced species of Trypetinae. thorax, which is mostly yellow with dark
Bigot (1880: 154, bottom of page) stated brown to black marks on the scutum, scu-
clearly that the species he described in this tellum, and subscutellum (Fig. 1; also see
paper were in his collection, but Adrian Freidberg and Kugler 1989, Figs. 188, 202,
Pont (personal communication) was unable 209). The microtrichia are pale or gray, ex-
to locate the single female (therefore holo- cept on some of the dark brown areas,
type) of S. caucasica in the Bigot Collec- which are bare or have dense, blackish mi-
tion at the University Museum, Oxford. He crotrichia. Except for the subscutellar
remarked that it is possibly there under marks, this pattern has been lost in C. in-
"some other name that Bigot changed to completa, which shares several synapomor-
caucasica with publication. He was prone phies with C. schineri and vesuviana (see
to doing this, and it causes endless confu- Fig. 5), indicating that it does belong in
sion." Carpomya (J. Jenkins, personal communi-
cation, has discovered a male genitalic syn-
Relationships and Classification
apomorphy that further supports this clade).
The Carpomyina was proposed as a sub- This color pattern is unique within the Car-
tribe of Trypetini by Norrbom (1989), but pomyina; in other genera, the thorax is dark
is currently considered a subtribe of Car- brown or black (except, in many species,
pomyini within the Trypetinae (Korneyev for the whitish postpronotal lobe, dorsal
1996). The desclerotized area at the apex of anepisternal stripe, apical part ofscutellum,
female syntergosternite 7 and the presence and sometimes a medial scutal spot or stripe)
of stomal guards, usually distinctly sclero- or predominantly yellow to orange, or if
tized, in the larva are probable synapomor- partly or mostly dark brown (e.g., Zonose-
phies for the subtribe (Norrbom 1989, Car- mata or Rhagoletis suavis species group),
roll 1992). The shape of the male surstyli the pattern is much different than in Car-
(inner and outer surstyli elongate, the latter pomya. Some species of Ceratitidina in the
with a long, apically directed posterior lobe Dacini do have thoracic color patterns sim-
and a short, mesally directed anterior lobe) ilar to Carpomya, thus convergent evolu-
C
is probably another synapomorphy of the tion of such a pattern in tica and the Old
Carpomyina or possibly for the Carpomyini World species of Carpomya is possible, but
(Korneyev 1996). Some reversal (reduction I have discovered no apomorphies shared
C
of the posterior lobe of the outer surstylus by tica and any other species of Carpo-
in a few taxa) must have occurred if this myina (or other Trypetinae) that would con-
character was in the groundplan ofthe Car- tradict the hypothesis that Carpomya, as de-
I
VOLUME 99, NUMBER 2 345
^(7.2)^ tica
=(5,6.1)H pardaUna
^(9,17,2)= •(8.2)==== wiedemanni
l!=(8.1,13)H (8.2)== schlneri
=
l!={6.2,7.1,15)H P=(5.0,6,3)= incompleta
=====
^(3,17.1): vesuviana
Fig. 5. Hypothesized phylogenetic relationships among species of Carpomya. Character numbers refer to
Table 1.
limited here, is monophyletic. Other char- (Bush 1966: 451), and I tentatively used the
acter state distributions indicate that the dis- R. tabellaria group, which has similar wing
tal location of r-m in C. tica and patterns, as the outgroup for analysis of
Cryptodacus and Haywardina cuculi is the character polarity.
result of convergence (see Norrbom The Hennig86 analysis of relationships
1994b). within Carpomya (based on the nine char-
Certain characters of the wing pattern acters with an asterisk in Table 1) resulted
may be additional synapomorphies of Car- in the single tree shown in Figure 5 (length
pomya, although some species now placed 17 steps, consistency index 82, retention in-
in Rhagoletis (particularly the tabellaria dex 75). Carpomya incompleta, schineri
species group) have similar wing patterns, and vesuviana form one monophyletic sub-
so some or all of these characters could be group, and C. pardalina and wiedemanni
plesiomorphic or synapomorphies at a high- form another. These groups together appear
er level within the Carpomyina. Better un- to be the sister group of C. tica. Despite
derstanding of the polarity and evolution of having several distinctive autapomorphies
wing patterns within the Carpomyina is (greatly elongated mouthparts and head, fa-
necessary to determine the phylogenetic cial carina shape, apically rounded first fla-
significance of these characters, but at least gellomere, and serrate aculeus tip; see
the strong similarity of the wing pattern of Freidberg and Kugler 1989, Figs. 201, 204),
C. tica with those of the other species of C. wiedemanni, the type species of Gonig-
Carpomya does not contradict its classifi- lossum, appears to be most closely related
cation in this genus. The similar elements to C. pardalina, which is the type species
of the wing pattern include: absence of the of Myiopardalis. The synonymy of Gonig-
accessory costal band; absence of the pos- lossum and Myiopardalis with Carpomya
terior apical band; the broad width of all of (type species C. vesuviana) is a subjective
the bands (reduced in C. incompleta) and decision ofranking, but their continued rec-
their yellow to brown color; discal and sub- ognition as monotypic genera (or the place-
apical bands transverse and parallel; and the ment of both species in Goniglossum)
at least partial separation ofthe anteriorapi- would require the proposal of yet another
C
cal band from the wing margin (Fig. 2; and monotypic genus for tica. Considering
Freidberg and Kugler 1989, PI. VIII, Figs. the diversity of species now classified in
4, 8, 10). Rhagoletis, I find the latter option undesir-
The relationship of Carpomya to other able. Already, Zaitzev (1919, 1947), Roh-
genera of Carpomyina has not been re- dendorf (1939) and Kandybina (1965,
solved. Its closest relatives may be within 1977) considered Myiopardalis a subgenus
the possibly paraphyletic genus Rhagoletis of Carpomya. The latter author found little
346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
difference among the larvae of the species Spilographa caucasica, and V. F. Zaitzev
here included in Carpomya, except that the arranged the loan of specimens from the
oral ridges are reportedly nonserrate in C. Zoological Institute, St. Petersburg. Alex-
wiedemanni. This character (not verified by ander Konstantinov and Raymond Gagne
C
personal examination by Kandybina in translated pertinent sections ofRussian and
wiedemanni) varies in other genera (e.g., French publications. John Jenkins and Am-
Anastrepha, see White and Elson-Harris non Freidberg kindly shared some unpub-
1992), and Kandybina otherwise considered lished data (a genitalic synapomorphy con-
C. wiedemanni larvae most similar to those firming the monophyly of the vesuviana
of C. pardalina and Rhagoletis flavigenu- group, and notes on the classification of
alis Hering of the tabellaria species group. Scleropithus, respectively), as well as their
thoughts regarding the relationships and
BlOGEOGRAPHY
classification of the Carpomyina. They, V.
The Carpomyina are distributed predom- Hernandez-Ortiz, G. J. Steck, and F. C.
inantly in the Holarctic Region and in high- Thompson reviewed earlier drafts of the
er altitude or temperate areas of the Neo- manuscript.
tropical Region. Although the Tephritidae
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