Thyreus species

ROZEN, JEROME G., 2000, Pupal Descriptions of Some Cleptoparasitic Bees (Apidae), with a Preliminary Generic Key to Pupae of Cleptoparasitic Bees (Apoidea), American Museum Novitates 3289, pp. 1-1 : 1-

publication ID	https://doi.org/ 10.1206/0003-0082(2000)289<0001:PDOSCB>2.0.CO;2
persistent identifier	https://treatment.plazi.org/id/A14D878F-FFE8-FFF5-829E-F9EBFD4F81D7
treatment provided by	Carolina
scientific name	Thyreus species
status

Treatment

Thyreus species ?

Figures 21–24 View Figs

DIAGNOSIS: The pupa of this species is similar to that of Xeromelecta californica , above, and is compared to other melectine

pupae in the diagnosis of the latter. Character states in boldface, below, differentiate the pupa of Thyreus species ? from that of X. californica .

HEAD: Integument without setae but with approximately three tubercles with sharply pointed, sclerotized apices on each side of upper frons (figs. 21, 24) (but not on vertex as in Xeromelecta californica ); scape without sharply pointed tubercles; rounded tubercles and verrucae absent. Apex of clypeus and labrum as described for X. californica except labrum as long as maximum

width; pupal ocelli well defined; most flagellomeres with welldefined swelling along ventral surface. Mandibles markedly swollen anteriorly and posteriorly, subapically; these swellings accommodating developing adult setae.

MESOSOMA: Integument spiculate in many areas; spicules especially long and conspicuous on mesoscutal tubercles (fig. 23); setae absent. Pronotum, mesepisternum, mesoscutal tubercles, and axillae as described for Xeromelecta californica except paired, spinebearing mesoscutal tubercles (figs. 21,23) much longer relative to basal width, with fewer spines, and nearly erect; mesoscutum without any other tubercles; mesoscutellar integument smooth, without spines; mesoscutellum with conspicuous pair of posteriorly directed tubercles overhanging metanotum (figs. 21, 24), corresponding to scutellar spines of adult; metanotum and tegula as described for X. californica . Mid and hind coxae with small, apically rounded tubercle on inside; trochanters without ventroapical projection; all femora without tubercles on outer surface.

METASOMA: As described for Xeromelecta californica except T1 without subapical row of very small, sharply pointed, mostly apically pigmented tubercles (figs. 21, 14). As in X. californica , apex of metasoma produced into short, broadly rounded projection as seen from above (fig. 22), this projection armed with small, median, sclerotized, thornlike spine and not with elongate, unsclerotized apical spine. Spiracles faintly evident.

MATERIAL STUDIED: One male pupa, S shore Issyk Kul, 10 km E KadshiSaj, Issyk Kul Region, Kyrgyzstan, 1675 m, 42°10'33̎ N 72°18'55 ̎E, VII51999 (J. G. Rozen) not in cocoon; from cell of Anthophora (Petalosternon) albifascies Alfken (kindly identified by Dr. Robert W. Brooks). Numerous adults (predominantly males) of this Thyreus were collected while patrolling the face of a vertical bank in which the host was nesting. These adults run to the distinctive ‘‘Group of Thyreus dimidiatapunctata (Spinola) ’’ in Lieftinck’s ( 1968) key to palearctic species and may represent an undescribed species near T. dimidiatipunctus or T. bidentatus (Kirby) .

REMARKS: Although adults of this species

lack maxillary palpi, the pupa possesses short, thin, distinct maxillary palpi (fig. 21).

GENERIC KEY TO PUPAE OF CLEPTOPARASITIC BEES

The following preliminary key is present ed from published accounts and from the information presented above. For completeness, the undescribed pupa of Coelioxoides (couplet 1) is included; it will be described in a forthcoming paper in which it will be compared with that of its host, Tetrapedia . Table 1 lists the species that provided the information for this key and references to published accounts.

1. Vertex, mesoscutum, and terga without hairs or tubercles; labrum longer than maximum width; pupa without cocoon. APIDAE (in part): TETRAPEDIINI...................... Coelioxoides Vertex, mesoscutum, and terga either with fine hairs or at least some of these structures with tubercles; labral length variable; pupa with or without cocoon ............................. 2

2(1). Vertex, mesoscutum, and terga with fine hairs, longest of which are at least four to five times length of minute tubercles from which they arise; 2 these tubercles so minute that terga appear to lack subapical, transverse row of tubercles; labrum elongate, much more than twice maximum diameter in frontal view; pupae found in cocoons. MEGACHILI DAE........................ 3 Vertex, mesoscutum, and terga without fine hairs, sometimes with sharply pointed tubercles whose apices may be setiform but these setiform apices nev er much more than length of tubercles from which they arise; these tubercles large enough that most terga appear to have subapical, transverse row of tubercles (figs. 1, 2, 5, 6); labrum usually about as long as maximum width or shorter, but, if longer ( Holcopasites , Oreopasites ), then vertex, mesoscutum, and terga with more or less conspicu

2 This character may not hold for Heterostelis , since Thorp ( 1966) commented that the pupa of H. hurdi was ‘‘apparently without long setae on’’ these surfaces. Nonetheless, fine hairs on these areas are common on most megachilid pupae, and it would be surprising, therefore, if they were not present.

ous sharp tubercles; pupae with or without cocoons............... 6

3(2). Axillae produced (correspond to axillae of adult) (Baker et al., 1985: fig. 6)........................ Coelioxys Axillae not produced............. 4

4(3). Mesoscutum not produced strongly posteriorly so that metanotum visible in dorsal view (Rozen, 1967: fig. 31); metanotum with single, median, round ed tubercle as seen from above (ibid.)........................ Dioxys Mesoscutum projecting posteriorly so that metanotum scarcely if at all visible in dorsal view (Rozen, 1966: fig. 18); metanotum without median tubercle... 5

5(4). Median section of mesoscutellum swollen, so that division of mesoscutum and mesoscutellum forming transverse groove as seen in lateral view (Rozen, 1966: fig. 17)......... Habrostelis Median section of mesoscutellum apparently not swollen, so that profile of mesoscutum and mesoscutellum nearly continuous in lateral view (Rust and Thorp, 1973: fig. 15)......... Stelis

6(2). Forewing with conspicuous tubercle midway to apex (Rozen, 1965: figs. 21, 22; Eickwort and Eickwort, 1972: fig. 7); subapical tubercles of terga rounded to acutely pointed but not bearing apical spine; hind tibia with large or small rounded tubercle at base; pupae without cocoons. HALICTIDAE 3 ..... 7 Forewing either without tubercle or, if present, tubercle low and not situated halfway to apex; subapical, tergal tubercles sharp pointed, not rounded, except those of Osiris and Isepeolus , which occur in cocoons; hind tibia without basal rounded tubercle. API DAE (in part)................. 8

7(6). Vertex with pair of large rounded tubercles in position of lateral ocelli, with pair of lower protuberances in front of upper end of compound eye, and with very low protuberance just above eye (Rozen, 1965: fig. 21).... Sphecodes Vertex with tubercles in position of lateral

3 As Dr. Michael S. Engel (personal commun.) has pointed out after reviewing the manuscript, pupae of only some parasitic Halictini are known; immatures of cleptoparasitic Augochlorini ( Temnosoma, Cleptommation , and Noctoraptor ) are unknown. However, the characters presented in this couplet correspond to those of the nonparasitic Augochlora pura (Say) (Michener, 1954) of the Augochlorini and therefore are likely to apply to the cleptoparasitic taxa of that tribe as well. ocelli small, very small tubercle just laterad of ocellar tubercle, without tubercle in front of upper end of eye or at summit of eye (Eickwort and Eickwort, 1972: fig. 7)....... Microsphecodes 4 8(6). Mesoscutum with single pair of conspicuous tubercles, each bearing numerous sharp spines (figs. 17, 18, 20, 21, 23, 24); upper frons and/or vertex with fine, sharply pointed tubercles (figs. 17, 18, 21, 24). MELECTINI.. Included taxa differentiated in diagnosis of Xeromelecta californica Mesoscutum with or without tubercles, but tubercles never bearing numerous spines; upper frons and vertex with or without sharply pointed tubercles.. 9 9(8). Tergal tubercles each with patch of dense spicules apically (figs. 7, 9, 10), without single, sharp point; mesoscutellum with pair of elevated, acutely pointed paramedian tubercles (fig. 17). OSIRI NI...................... Osiris Most tergal tubercles each with single, short or attenuated, sharp apex, always without apical patch of dense spicules; mesoscutellum with or without pair of tubercles, but tubercles apically round ed when present (figs. 11, 15)... 10 10(9). Labral apex bilobed as seen in frontal view (fig. 11)................ 11 Labral apex truncated, curved, or broadly pointed, not bilobed in frontal view............................ 13 11(10). Mesoscutum without pair of rounded tubercles (Camargo et al. (1975: fig. 11); midtibial spur normal, tapering to apex. RHATHYMINI........ Rhathymus Mesoscutum with pair of rounded tubercles (fig. 11); midtibial spur truncate or bilobed (fig. 11), not tapering to point ed apex. ERICROCIDINI....... 12 12(11). Tegula with pronounced tubercle (Rozen and Buchmann, 1990: fig. 61)....

....................... Ericrocis Tegula only slightly produced, without distinct tubercle (fig. 11)... Mesoplia 13(10). Tergal tubercles apparently without apically pointed tip under normal magnification but larger tubercles actually with tiny dotlike spine under high magnification (fig. 15); axillae strongly produced (fig. 15); tubercles in position of

4 Although Eickwort and Eickwort ( 1972) pointed out that Microsphecodes kathleenae is a social parasite, oth er species in the genus may be cleptoparasites, and hence the genus is included.

lateral ocelli pronounced, rounded, without sharp apices (fig. 15). ISEPEO LINI.................. Isepeolus Most tergal tubercles with distinct, sharply pointed apices under normal magnification; axillae usually not produced, but, if so (some Nomadinae), tubercles of head with pointed apices..... 14

14(13). Tegula without distinct tubercle and vertex without sharply pointed tubercles; mesoscutum with pair of tubercles that are rounded or apically verrucose and located in line with or in front of tegula in lateral view (Rozen et al., 1978: fig. 34; RoigAlsina and Rozen, 1994: figs. 20, 24); pupa found in cocoon. PRO TEPEOLINI........... Leiopodus Either tegular tubercle well developed (Rozen, 1977: figs. 21, 26; 1994: fig. 20; 1997b: fig. 25) or vertex with sharply pointed tubercles; mesoscutum with or without tubercles; if present, tubercles either sharply pointed or (in Brachynomada ) somewhat posterior to tegula in lateral view (Rozen, 1994: fig. 20; 1997b: fig. 25); pupa without cocoon. NOMADINAE.......... 15

15(14). Tegula with conspicuous tubercle (Rozen, 1977: figs. 21, 26; 1994: fig. 20; 1997b: fig. 25); vertex, mesoscutum, mesoscutellum without sharply pointed tubercles (see same figures); sterna without tubercles. BRACHYNOMADINI ............................ 16 Tegula without tubercle (figs. 1–4, 5, 6); usually vertex and often mesoscutum and mesoscutellum with sharply point ed tubercles (figs. 1, 2, 5, 6); if vertex without sharp tubercles ( Odyneropsis ), some sterna with apical row of sharp tubercles.................... 18

16(15). Mesoscutum with pair of small, rounded tubercles (Rozen, 1994: fig. 20)....................... Brachynomada Mesoscutum without tubercles (Rozen, 1977: figs. 21, 26)............. 17

17(16). Forewing with distinct, acutely rounded tubercle along anterior margin, approximately twothirds way to apex (Rozen, 1977: fig. 26); vertex with tubercles somewhat more pronounced.......................... Melanomada Forewing with only vague swelling along anterior margin approximately twothirds way to apex (Rozen, 1977: fig. 21); vertex with tubercles vaguely expressed.............. Paranomada

18(15). Apical lateral angle of clypeus with downwarddirected tubercle (Rozen and McGinley, 1974: fig. 29); sterna without apical, sharply pointed tubercles. AMMOBATINI.......... 19 Apical lateral angle of clypeus normal, without tubercle; some sterna with apical, sharply pointed tubercles (fig. 6) (these tubercles very small and easily overlooked in Neolarra , fig. 1)... 20 19(18). Terminal metasomal spine moderately long (Rozen and McGinley, 1974: fig. 26); mesoscutellum with sharply point ed tubercles (ibid.; Rozen, 1992: figs. 5, 8)................ Oreopasites Terminal metasomal spine small, short (Rozen and McGinley, 1974: figs. 32, 33); mesoscutellum without sharply pointed tubercles (ibid.).... Pasites 20(18). Tubercles of vertex and most metasomal terga with apices produced as straight or curved, elongate hair, often as long a tubercle itself (fig. 1). NEOLARRINI ....................... Neolarra Tubercles of vertex and metasomal terga with sharp apices short, not hairlike; if somewhat elongate ( Holcopasites ), then apices hooked or zigzagged (figs. 5, 6)........................ 20 20(19). Terminal spine not developed (Rozen, 1989: fig. 24), so that metasoma ending in short triangular projection in dorsal view; axillae often strongly projecting posteriorly. EPEOLINI......... 21 Terminal spine elongate (figs. 1, 6), about as broad as high; axillae not strongly projecting posteriorly (figs. 2, 5).. 22 21(20). Vertex and mesoscutum 5 apparently without sharply pointed tubercles.....

.................... Odyneropsis Vertex and mesoscutum with numerous sharply pointed tubercles........

.............. Epeolus , Triepeolus 22(20). Labrum elongate, much longer than maximum width (labrum with basal swelling, fig. 6, that should not be confused with apex that is posterior to closed mandibles); tubercles of head and elsewhere with long apices some of which are kinked or zigzag. AMMOBATO IDINI............... Holcopasites Labrum somewhat shorter than maximum width, its apex just reaching closed mandibles (Rozen, 1997a: fig. 1); tubercles of head with apices that are at most hooked. BIASTINI..... Neopasites

5 Evaluated on the basis of a single, poorly preserved specimen.

TABLE 1

Taxa Used for the Generic Key to Pupae of Cleptoparasitic Bees

(References are to published descriptions of material examined. Early references taken in large part from the work of McGinley, 1989. AMNH = American Museum of Natural History.)

HALICTIDAE

Sphecodes albilabris (Kirby) Rozen ( 1965) View in CoL

Microsphecodes kathleenae (Eickwort) Eickwort & Eickwort ( 1972) View in CoL

MEGACHILIDAE View in CoL

Stelis cholorocyanea (Cockerell) Heterostelis hurdi Thorp

Hoplostelis bilineolata (Spinola) Dioxys pomonae pomonae (Cresson) Coelioxys sayi Robertson C. lativentris Friese

Rust & Thorp ( 1973) Thorp ( 1966)

Rozen ( 1966)

Rozen ( 1967)

Baker et al. ( 1985) In AMNH collection

APINAE

Ericrocis lara (Cresson) Rozen & Buchmann ( 1990)