Corresponding authors: Charuwat Taekul (
Academic editor: Michael J. Sharkey
The genus
Species of
The known hosts of species in the subfamilies
In more than a century since its original description,
This work is based upon specimens in the following collections, with abbreviations used in the text: AEIC, American Entomological Institute, Gainesville, FL
Abbreviations and morphological terms used in text: A1, A2, ... A12: antennomere 1, 2, … 12; claval formula: distribution of the large, multiporous basiconic sensilla on the underside of apical antennomeres of the female, with the segment interval specified followed by the number of sensilla per segment (
In the Material Examined the numbers prefixed with “OSUC” are unique identifiers for the individual specimens. The label data for all specimens have been georeferenced and recorded in the
Data associated with the genus
In this article we have followed the precedent of
This work is conducted as part of the
Moderate-sized, length 3.0–5.6 mm; head prognathous, flattened anteroposteriorly, mesosoma and metasoma strongly dorsoventrally depressed; body black; macropterous.
Head in dorsal view strongly transverse; vertex laterad of posterior ocellus smooth or with few faint striae, between posterior ocelli finely longitudinally striate; hyperoccipital carina present as fine ridge on vertex between compound eyes; occipital carina absent; posterior ocellus distinctly separated from inner orbit of compound eyes, OOL > diameter of ocellus; compound eye large, appearing glabrous; frons without depression, shallowly convex, with median longitudinal sulcus bifurcating dorsally near median ocellus and ventrally near toruli; interantennal process well-developed, narrow; torular triangle present; submedian carina sometimes present; orbital carina sometimes present; lower frons, including cheek, with weak fanlike striae arising from mandibular condyle; shortest distance on frons between eyes less than eye height; inner orbits weakly diverging ventrally; postclypeus strongly projecting above anteclypeus, subtriangular, anteclypeus short, longest medially, lateral corners not produced; malar sulcus present; gena variably expanded, smooth to longitudinally striate or with few faint striae; labrum hidden by clypeus; mandible short, apex tridentate, middle tooth distinctly shortest, teeth arranged transversely; maxillary palpus 2-segmented, all segments cylindrical; labial palpus 1-segmented, very short; antenna 12-merous in both sexes; radicle very broad, inserted into ventral apex of A1, nearly parallel to longitudinal axis of A1, with small lateral spine; A1 almost triangular and expanded outwardly into spine, particularly in female; A2 distinctly shorter than A3; gustatory sensilla on female antenna arranged in longitudinal pairs on apical antennomeres; clava laterally compressed, claval formula A12–A8: 1-2-2-2-1; male antenna with tyloid on A5.
Mesosoma in dorsal view longer than wide, in lateral view much longer than high; pronotum in dorsal view strongly narrowed laterally, anterolateral corner weakly angulate; transverse pronotal carina absent; vertical epomial carina absent; dorsal epomial carina absent; lateral face of pronotum concave; netrion absent; anterior margin of mesoscutum strongly flexed ventrally to meet pronotum; mesoscutum semicircular in outline, posterolateral corner rounded; parapsidal line variably developed; notaulus variably developed: absent, nearly percurrent or percurrent, mesoscutum sometimes with strong sublateral carina paralleling notaulus; skaphion absent; prespecular sulcus and posterior mesepimeral sulcus present; speculum smooth, rarely longitudinally striate (
Metasoma elongate, parallel-sided, strongly flattened dorsoventrally; female with six visible terga and sterna, male with seven visible terga and sterna; second to fifth segments equal in length, third and fourth widest, subequal to each other in size; submarginal ridge developed, defined by narrow laterotergites to form deep submarginal rim; sublateral carina well developed on tergites, rarely absent (
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Striae within ocellar triangle sparse (fewer than 20); distance between anterior ocellus and posterior ocellar line in frontal view greater than or equal to 0.5× POL (Figs. |
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- | Striae within ocellar triangle dense (more than 20); distance between anterior ocellus and posterior ocellar line in frontal view less than 0.5× POL (Figs. |
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Sculpture on T1 longitudinally striate, interstices with coriaceous microsculpture (Fig. |
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- | Sculpture on T1 longitudinally striate throughout (Fig. |
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Frontal sculpture between inner orbit and central keel longitudinally striate, striae either extending through most of length of frons, or with few (4–5) striae restricted to upper half of frons (Figs. |
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- | Frons between inner orbit and central keel smooth (Figs. |
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Mesoscutum with one lateral carina; notaulus absent (Fig. |
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- | Mesoscutum with two lateral carinae; notaulus present (Fig. |
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Posterior scutellar sulcus complete (Figs. |
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- | Posterior scutellar sulcus interrupted medially (Figs. |
Body length of male: 2.95–4.35 mm (n=20). Body length of female: 3.46–4.28 mm (n=20).
Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: densely striate. Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Orbital carina: absent. Sculpture of malar region: smooth, faintly longitudinal striae with limited microsculpture near eye.
Color of female antenna: A1–A7 yellow to light brown, A8–A12 dark brown to black. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: yellow to light brown throughout.
Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even. Notaulus: present. Notaulus form: mesal and lateral margin arched. Length of notaulus: abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus anteriorly: parallel-sided. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: 0. Medial carina of mesoscutum: absent. Parapsidal line: present. Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparse (less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: longitudinally striate or with few reticulations. Median propodeal sulcus: widened posteriorly. Sculpture of submedian propodeal field: longitudinally striate. Posterolateral margin of propodeum: longitudinally striate laterally, rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise yellow throughout, rarely hind femur dark brown.
Female Postmarginal vein: absent. Fore wing: hyaline.
Sculpture on T1: longitudinally striate with setigerous punctures medially. Sublateral carina on T2–T4: present anteriorly, absent posteriorly. Sculpture on T2–T4: setigerous punctures throughout, longitudinally striate anteriorly.
Holotype female: CAMEROON: Garoua (deposited in MNHN). Other material: (78 females, 57 males) BENIN: 14 females, 5 males, CASENT 2137991 (CASC); OSUC 207951-207968 (CNCI). CAMEROON: 17 females, 7 males, BMNH(E)#790211, 848520-848536 (BMNH); CASENT 2137986 (CASC); OSUC 250657-250661 (CNCI). CENTRAL AFRICAN REPUBLIC: 1 female, 1 male, OSUC 176086, 247778 (SAMC). GHANA: 2 males, BMNH(E)#790201, 848510 (BMNH). GUINEA: 1 female, 1 male, OSUC 207895, 250625 (CNCI). GUINEA-BISSAU: 1 female, OSUC 253728 (MZLU). IVORY COAST: 22 females, 13 males, OSUC 207977-207983, 207986-208007 (CNCI); OSUC 58731-58736 (OSUC). KENYA: 2 females, 7 males, OSUC 173854, 207969-207976 (CNCI). MOZAMBIQUE: 3 males, OSUC 207948-207950 (CNCI). NIGERIA: 9 females, 5 males, BMNH(E)#790200 (BMNH); OSUC 250639-250650 (CNCI); OSUC 173856 (OSUC). SIERRA LEONE: 2 females, 5 males, BMNH(E)#790195, 848506 (BMNH); OSUC 253722-253726 (MZLU). SOUTH AFRICA: 2 females, 5 males, OSUC 207938, 207943-207946, 250663 (CNCI); OSUC 253727 (MZLU). TANZANIA: 3 females, 1 male, OSUC 253741-253744 (SAMC). TOGO: 1 female, OSUC 253754 (CNCI). UGANDA: 1 female, 1 male, OSUC 207984-207985 (CNCI). YEMEN: 1 female, 1 male, OSUC 250651-250652 (CNCI). ZIMBABWE: 1 female, BMNH(E)#790209 (BMNH).
This species is widespread in the Afrotropical realm, extending from east Africa to Yemen and south to the north of South Africa. The color of the female antenna is variable: the scape is yellow to light brown, but in some specimens is dark brown to black (OSUC 207985, 207972;
Body length of male: 3.87 mm (n=1). Body length of female: 3.48–3.97 mm (n=3).
Length between anterior ocellus and posterior ocellar line in frontal view: greater than or equal to 0.5 times POL. Striae within ocellar triangle: sparse (equal to or less than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.
Color of female antenna: dark brown to black throughout. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: dark brown to black throughout.
Sculpture on medial lobe of mesoscutum: longitudinally striate. Setation of medial lobe of mesoscutum: sparse to glabrous. Notaulus: present. Notaulus form: mesal margin arched, lateral margin straight. Length of notaulus: percurrent or nearly so. Width of notaulus anteriorly: narrowed anteriorly. Pilosity of notaulus: present. Number of lateral carinae on mesoscutum: 0. Medial carina of mesoscutum: present. Parapsidal line: present. Posterior scutellar sulcus: interrupted medially or complete. Setation of posterior half of ventral metapleural area: sparse (less than 25 setae). Metascutellum size: narrow, metanotum lateral to metascutellum with 4–6 foveae. Sculpture on ventral metapleural area: longitudinally striate, coriaceous microsculpture within interstices or strongly reticulate rugose with foveolae. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: longitudinally striate, in male interstices with coriaceous microsculpture. Posterolateral margin of propodeum: reticulate rugose with foveolae throughout. Color of legs: coxae dark brown to black, otherwise yellow throughout.
Postmarginal vein: reduced, stumplike. Fore wing: hyaline.
Sculpture on T1: longitudinally striate, coriaceous microsculpture within interstices. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: longitudinally striate with coriaceous microsculpture within interstices.
The epithet arcuatus, Latin for bent like a bow, refers to the shape of the notaulus.
Holotype female: AUSTRALIA: Western Australia, Keystone Rd., 3 km W Walpole, 34°59.01'S 116°40.76'E, George, Hawks, Munro, YPT, OSUC 250635 (deposited in ANIC). Paratypes: AUSTRALIA: 3 females, 1 male, OSUC 250633–250636 (CNCI).
Body length of male: 3.46–4.20 mm (n=7). Body length of female: 3.14–4.24 mm (n=8).
Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: densely striate. Frontal sculpture between inner orbit and central keel: longitudinally striate, striae extending through most of length of frons. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Orbital carina: present. Sculpture of malar region: longitudinally striate or with few faint striae.
Color of female antenna: A1–A7 yellow to light brown, A8–A12 dark brown to black. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: brown or dark brown to black throughout.
Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even. Notaulus: absent. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: 1. Medial carina of mesoscutum: absent. Parapsidal line: present. Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparse (less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: smooth anteriorly, coarsely foveolate punctate posteriorly. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: smooth throughout or with few faint striae. Posterolateral margin of propodeum: smooth laterally, longitudinally striate to rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise variable.
Postmarginal vein: absent. Fore wing: hyaline.
Sculpture on T1: longitudinally striate laterally, uniformly setigerous punctate medially. Sublateral carina on T2–T4: present anteriorly, absent posteriorly. Sculpture on T2–T4: setigerous punctures throughout, longitudinally striate anteriorly.
The epithet mysterium, Latin for mystery, refers to the interpretation of the mesoscutal carinae.
Holotype female: BOTSWANA: Serowe, Farmer’s Brigade, 22°22.998'S, 026°43.002'E, May 1989, Malaise trap, P. Forchhammer, OSUC 171372 (deposited in USNM). Paratypes: (11 females, 14 males) BOTSWANA: 6 females, CASENT 2137987-2137990 (CASC); OSUC 250665 (CNCI). SOUTH AFRICA: 3 females, 13 males, BMNH(E)#790187-790189, 790196-790197, 790199, 848507-848509 (BMNH); OSUC 207935-207937, 207939, 207942, 207947 (CNCI); OSUC 230254 (OSUC). ZIMBABWE: 3 females, 1 male, OSUC 250653-250656 (CNCI).
Some specimens show variability in the prominence of the sculpture between inner orbit and central keel on the frons.
Body length of male: 3.24–4.71 mm (n=20). Body length of female: 3.31–5.59 mm (n=20).
Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Orbital carina: absent. Sculpture of malar region: smooth, longitudinally striate or with few faint striae. Color of female antenna: dark brown to black throughout, A1–A7 yellow to light brown, A8–A12 dark brown to black, or A1–A7 brown, A8–A12 dark brown to black, in some antennae sequentially darker from scape to apex. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: brown.
Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even. Notaulus: present. Notaulus form: mesal and lateral margin arched. Length of notaulus: percurrent or nearly so, or abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus anteriorly: parallel-sided. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: 0. Medial carina of mesoscutum: absent. Parapsidal line: present or absent or faint. Posterior scutellar sulcus: interrupted medially. Setation of posterior half of ventral metapleural area: dense (more than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: smooth to faintly longitudinally striate, or longitudinally striate or with few reticulations. Median propodeal sulcus: narrow throughout length, widened posteriorly. Sculpture of submedian propodeal field: smooth throughout or with few faint striae or longitudinally striate. Posterolateral margin of propodeum: smooth laterally, longitudinally striate to rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise brown, or coxae dark brown to black, otherwise yellow throughout.
Postmarginal vein: absent. Fore wing: hyaline, infuscate.
Sculpture on T1: longitudinally striate laterally, uniformly setigerous punctate medially. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: uniformly setigerous punctate.
Holotype female,
Other material: (121 females, 71 males, 6 unknowns) AUSTRALIA: 1 female, 4 males, BMNH(E)#790202-790203, 790208 (BMNH); OSUC 250637 (CNCI); OSUC 141949 (OSUC). BANGLADESH: 1 female, OSUC 173855 (CNCI). CHINA: 2 females, 4 males, BMNH(E)#848537-848540 (BMNH); OSUC 321841-321842 (CNCI). GUAM: 3 females, 1 male, 3 unknowns, OSUC 250638 (CNCI); OSUC 207850-207852, 207856-207858 (USNM). INDIA: 7 females, 2 males, BMNH(E)#790185, 790206-790207, 848518-848519 (BMNH); OSUC 230648 (OSUC); OSUC 207838-207840 (USNM). INDONESIA: 13 females, 9 males, BMNH(E)#790210 (BMNH); OSUC 207859-207874, 207876, 207893-207894, 207896 (CNCI); OSUC 204851 (UCDC). JAPAN: 1 female, OSUC 173082 (UASK). MALAYSIA: 9 females, 5 males, BMNH(E)#790193, 790204 (BMNH); OSUC 207877-207883, 207885-207887, 207892 (CNCI); OSUC 207884 (SCAU). PAPUA NEW GUINEA: 24 females, 21 males, BMNH(E)#790205, 848511-848517 (BMNH); CASENT 2137992-2138022 (CASC); OSUC 207897, 250626-250628 (CNCI); OSUC 160037-160038 (EMEC). PHILIPPINES: 3 females, 7 males, BMNH(E)#790184 (BMNH); OSUC 207875 (CNCI); OSUC 207842-207849 (USNM). SOLOMON ISLANDS: 1 female, BMNH(E)#848543 (BMNH). TAIWAN: 18 females, 11 males, OSUC 173803-173831 (TARI). THAILAND: 36 females, 2 males, 2 unknowns, BMNH(E)#790190, 790192, 790194, 848502-848505, 848541-848542 (BMNH); OSUC 207888-207890 (CNCI); OSUC 207837, 215797, 253701-253721, 253760-253762, 253764 (OSUC); OSUC 207891 (SCAU). VANUATU: 2 males, 1 unknown, BMNH(E)#790183, 790186, 790198 (BMNH). VIETNAM: 2 females, 3 males, OSUC 277708-277709 (RMNH); OSUC 184378-184380 (ZMAS).
Some specimens show variability in antenna and leg color. Because of the uniformity of the principal characters – notaulus form and length, interrupted posterior scutellar sulcus, and the propodeal sculpture – we consider these specimens to be conspecific with
Body length of male: 3.05–3.22 mm (n=2). Body length of female: 2.89–3.40 mm (n=2).
Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: with few (4–5) striae, striae limited to upper half of frons. Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.
Color of female antenna: A1–A7 brown, A8–A12 dark brown to black, in some antennae sequentially darker from scape to apex. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 straight, antennomeres distinctly separated. Color of male antenna: dark brown to black throughout.
Sculpture on medial lobe of mesoscutum: longitudinally striate. Setation of medial lobe of mesoscutum: sparse to glabrous. Notaulus: present. Notaulus form: mesal margin arched, lateral margin straight. Length of notaulus: abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus anteriorly: narrowed anteriorly. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: 2. Medial carina of mesoscutum: absent. Parapsidal line: absent or faint. Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparse (less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: strongly reticulate rugose with foveolae. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: smooth throughout or with few faint striae. Posterolateral margin of propodeum: longitudinally striate laterally, rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise yellow in female and brown in male.
Postmarginal vein: absent. Fore wing: hyaline.
Sculpture on T1: longitudinally striate throughout. Sublateral carina on T2–T4: absent. Sculpture on T2–T4: longitudinally striate throughout, uniformly setigerous punctate within interstices on T3–T4.
The epithet striga, Latin for swath, refers to the distinct striae of the frons.
Holotype female: AUSTRALIA: Western Australia, Cape Arid N.P., 33°31.134'S, 123°26.052'E (WDPA-UN), 30.XII–3.I.1987, Malaise trap, J.S. Noyes, OSUC 250630 (deposited in ANIC). Paratypes: AUSTRALIA: 1 female, 2 males, OSUC 250629, 250631, 250632 (CNCI).
Body length of male: 3.21–4.53 mm (n=20). Body length of female: 3.14–4.23 mm (n=20).
Length between anterior ocellus and posterior ocellar line in frontal view: greater than or equal to 0.5 times POL. Striae within ocellar triangle: sparse (equal to or less than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.
Color of female antenna: dark brown to black throughout. Female outer lateral apex of scape: bluntly rounded. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: dark brown to black throughout.
Sculpture on medial lobe of mesoscutum: longitudinally striate. Setation of medial lobe of mesoscutum: moderately dense, even. Notaulus: present. Notaulus form: mesal and lateral margin arched. Length of notaulus: percurrent or nearly so. Width of notaulus anteriorly: parallel-sided. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: 0. Medial carina of mesoscutum: absent. Parapsidal line: present or absent or faint. Posterior scutellar sulcus: interrupted medially. Setation of posterior half of ventral metapleural area: sparse (less than 25 setae). Metascutellum size: narrow, metanotum lateral to metascutellum with 4–6 foveae. Sculpture on ventral metapleural area: longitudinally striate or with few reticulations. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: longitudinally striate. Posterolateral margin of propodeum: margined by coarsely punctate furrow. Color of legs: coxae dark brown to black, otherwise brown.
Postmarginal vein: reduced, stumplike. Fore wing: hyaline or infuscate.
Sculpture on T1: longitudinally striate throughout. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: T2 longitudinally striate throughout, T3–T4 longitudinally reticulate laterally, smooth medially.
The epithet mzantsi, Xhosa for south, is a reference to the the collecting locality.
Holotype female: SOUTH AFRICA: 34°27.414'S 19°21.393'E Western Cape, Walker Bay Nat. Res., 57 m, Site1, Malaise trap, S. Coast Strandveld, 4.X–1.XI.1997, S.van Noort, WB97–M11, OSUC 243790 (deposited in SAMC). Paratypes: SOUTH AFRICA: 63 females, 31 males, OSUC 202440–202441 (AEIC); OSUC 207908–207934, 207941, 250662, 250664 (CNCI); OSUC 266101–266102 (MZLU); OSUC 188488–188489, 207830–207836, 207898–207907, 207940, 213995, 226020–226023, 237213–237217, 243454–243458, 243506–243507, 243790–243791, 253729–253740, 253745–253753 (SAMC).
Thanks to A. Bennett, M. Buffington, R. Danielsson, J. LaSalle, S. Myers, D. Notton, T. Nuhn, W. Pulawski, S. van Noort, L. Vilhelmsen, C. Villemant, and R. Zuparko for access to and loans of material for this study; A. Mayekiso for consultation on Xhosa; and to L. Musetti, A.D. Austin, M. Yoder for much valued assistance in this work. This material is based upon work supported in part by the National Science Foundation under grant No. DEB–0614764. C.T. was supported by a fellowship from the Agricultural Research Development Agency, Ministry of Agriculture and Cooperatives, Thailand.