TWO NEW SOUTHERN AFRICAN APATOMYZA WIEDEMANN ( DIPTERA , BOMBYLIIDAE , CROCIDIINAE ) WITH DISCUSSION ON THEIR PHYLOGENETIC POSITION

Two new species of Apatomyza Wiedemann from South Africa, A. whocantell spec. nov. and A. angusticephala spec. nov., are described, illustrated, and placed within the most recent key to species. Into the data matrix recently used to infer a phylogenetic hypothesis for the subfamily, were added the two new taxa, in order to verify their position and relationships. They form the most apical clade in Apatomyza.


INTRODUCTION
The Afrotropical genus Apatomyza was erected by Wiedemann (1820) to include a single species: A. punctipennis.Lamas et al. (2001) revised Apatomyza and proposed Adelogenys Hesse as a junior synonym.They also described three new species (A.capensis, A. desertica and A. irwini) which, together with the type species A. punctipennis and the previous species included in Adelogenys by Hesse (1938) (A. braunsii, A. culicoides and A. namaquensis -junior synonym of A. punctipennis), comprised a total of six species in the genus.Lamas et al. (2003) described four new Apatomyza (A. bethuliensis, A. proboscidea, A. aurea and A. glabripleura) and presented an identification key to all included species.Evenhuis & Greathead (2003), following the taxonomic changes proposed by Lamas et al. (2001Lamas et al. ( , 2003)), considered ten species among Apatomyza: A. aurea Lamas & Couri (2005) proposed a phylogenetic hypothesis for the Crocidiinae in which the subfamily was divided in two tribes (Desmatomyiini and Crocidiini), and Apatomyza was shown to be a monophyletic clade within the Crocidiini.
Studying a new series of unidentified material from the Natal Museum (Pietermaritzburg, South Africa), the authors found two new Apatomyza species from Southern Africa, which are herein described and illustrated.Their position in the key of Lamas et al. (2003) and in the phylogenetic hypothesis of Lamas & Couri (2005) are given.

Cladistic Analysis
Cladistic analysis was carried out using parsimony as implemented in Nona (Goloboff, 1993) version 2.0, running in Winclada (Nixon, 2002) version 1.00.08.The two new described taxa (Apatomyza whocantell spec.nov.and A. angusticephala spec.nov.) were added to the Crocidiinae data matrix of Lamas & Couri (2005), with 30 morphological characters, in order to verify their position.The cladograms were generated, as in Lamas & Couri (2005), applying successive weighting (with the macro swt.run associated to NONA) and with the characters 24, 27, 28 and 29 treated as non-additive.In an attempt to quantify the support for the clades, we have calculated the Bremer support (Bremer, 1994) for each node with Nona (Goloboff, 1993) using equally weighted characters: hold 10000, hold*, suboptimal 30, mult*1000.Branch support was also verified by using Jacknifing.The lines included in the matrix are shown in Appendix 1.

RESULTS
Apatomyza whocantell spec.nov.Diagnosis: A. whocantell spec.nov. is morphologically closed to A. proboscidea Lamas, Evenhuis & Couri, 2003, of which it can be easily distinguished by the infuscation of the wings and the color pattern of the mesonotum.
Male: Body length: 4.0 mm; wing length: 3.9 mm.Head: eyes separated at level of anterior ocellus by about two times width of ocellar tubercle; frons gray pollinose with dark brown bristles; face gray pollinose; oral-genal cup polished dark brown with a yellow mark laterally; first palpal segment dark brown, 1/3 of proboscis length, with short light brown hairs; second palpal segment oval, length about 1/4 of first segment; proboscis dark brown; labellum about 1/5 of proboscis length; antenna with scape dark brown, gray pollinose, 2 x longer than pedicel, with dark brown bristles; flagellum dark brown, 4 x longer than pedicel, with two flagellomeres and a short hyaline apical style; ocellar tubercle gray pollinose with dark brown bristles; occiput gray pollinose with dark brown bristles, white scales and yellow hairs.
Thorax: mesonotum (Fig. 1) gray pollinose, with sparse short dark brown bristles and yellow scales, with paired admedian dark brown stripes, and an additional lateral stripe; scutellum gray pollinose, with long yellow bristles, yellow scales and short dark brown bristles; postpronotal lobe with long yellow hairs; prealar and supra-alar bristles yellow; postalar callus with dark brown bristles; pleura gray pollinose; anepisternum with white scales and long yellow hairs on upper portion; katepisternum with white scales and hairs centrally; anepimeron, meropleurite, laterotergite, mediotergite, metepisternum and metepimeron bare; halter with stem and knob yellow, with brown marks and yellow hairs ventrally.
Legs: Coxae concolorous with pleura, with white scales and sparse white hairs; femora dark brown, gray pollinose, with white scales on anterior, dorsal and posterior surface and white hairs on ventral surface; tibiae yellow with yellow bristles; tarsi yellow; claws light brown on base and dark brown on tip; pulvilli and claws similar in length.
Wing (Fig. 2): hyaline, slightly infuscated, with light brown veins and dark brown areas: on humeral vein; base of r2+3, center of br and base of r1; apex of bm, base of discal and upper center of cup; center of r1, r2+3, around r-m and center of dm; lower center of dm, base of m2 and center of cua1; surrounding m-m; base of r4 and margin of r2+3; r-m crossvein beyond middle of discal cell but before end of vein Sc. Discal cell narrowed apically; anal cell closed before wing margin; squama hyaline with yellowish white fringe.
Abdomen: dark brown, with sparse yellowish white hairs, longer hairs on lateral margins.
pump, uniting to form a short common spermathecal duct; sperm pump placed in the middle 1/3 of the spermathecal duct; sclerotized collars developed on both ends of sperm pumps; spermathecal duct with the same width throughout.

Notes:
The unique male holotype is pinned in copula with a female paratype.In order to preserve this interesting mount (Fig. 4) the male terminalia were not dissected.
Etymology: named for the farm "Who can Tell" (Cradock, Eastern Cape) on which the type series was collected.The species epithet is treated here as a noun in apposition.Apatomyza angusticephala spec.nov.Diagnosis: A. angusticephala spec.nov.and A. whocantell spec.nov., key out near A. proboscidea Lamas, Evenhuis &Couri, 2003 andA. bethuliensis Lamas, Evenhuis &Couri, 2003, in the key of Lamas et al. (2003), as they have the second palpal segment oval.The length of proboscis indicates a close relation between A. angusticephala spec.nov.with A. bethuliensis (short proboscis) and A. whocantell spec.nov.with A. proboscidea (long proboscis). A. angusticephala spec.nov.could be easily segregated from A. bethuliensis by the dorsoventrally flattened head and color pattern of mesonotum.
Head: slightly dorsoventrally flattened (Fig. 5); eyes separated at level of anterior ocellus for about the width of ocellar tubercle; frons and face gray pollinose; oralgenal cup polished dark brown with a yellow mark laterally; first palpal segment dark brown, 1/2 of proboscis length, with short light brown hairs; second palpal segment oval, length about 1/5 of first segment; proboscis dark brown; labellum about 1/3 of proboscis length; antenna with scape dark brown gray pollinose, 3 x longer than pedicel, with dark brown bristles; flagellum dark brown, 5 x longer than pedicel, with three flagellomeres and a short hyaline apical style; ocellar tubercle gray pollinose with dark brown bristles; occiput gray pollinose with dark brown bristles.
Thorax: mesonotum (Fig. 6) dark brown, with a median longitudinal gray pollinose stripe, and two admedian dark brown stripes, with sparse short dark brown bristles; scutellum polished brown, gray pollinose, with long dark brown bristles; postpronotal lobe with long dark brown hairs; prealar and supra-alar bristles dark brown; postalar callus with dark brown bristles; pleura gray pollinose; anepisternum with sparse white hairs on upper portion; katepisternum with sparse white hairs; anepimeron, meropleurite, laterotergite, mediotergite, metepisternum and metepimeron bare; halter with stem light brown and a well developed yellow knob.
Legs: Coxae concolorous with pleura, with white hairs on anterior surface; femora dark brown, gray pollinose, with white hairs and scales; tibiae and tarsi dark brown with dark brown bristles; claws light brown on base and dark brown on tip; pulvilli and claws similar in length.
Female: as in male, except for: Body length: 2.5 mm; wing length: 2.3 mm.Eyes separated at level of anterior ocellus for about 3 x width of ocellar tubercle.
Spermathecae (Fig. 9): spermathecal reservoir pear shaped with longitudinal axis equal to sperm pump length; spermathecal duct 4 times longer than sperm pump, uniting to form a short common spermathecal duct and with a sclerotized moniliform pattern apically, fused with the spermathecal reservoir; sperm pump

Identification key
Both new species were added to the key of Lamas et al. (2003)

DISCUSSION
The two new Apatomyza species are easily segregated from the 10 previously known.Despite some morphological characters indicating similarities with A. proboscidea and A. bethuliensis, this relation is not corroborated by cladistic analysis.The tree obtained by successive weighting (Fig. 10) has 81 steps of length with a consistency index of 40 and a retention index of 74, and presents the same topology as the tree presented by Lamas & Couri (2005) except for the two new species included herein.The two new taxa (A. angusticephala spec. nov. and A. whocantell spec. nov.) appeared forming an apical clade with A. braunsii as its sister group (Fig. 10).This clade is supported by the reversion of character 9 (micropubescence in the laterotergite), where the apomorphic condition (presence of micropubescence) is a synapomorphy of the entire Crocidiini, and the homoplastic character 27, where the apomorphic condition is shared with the clade formed by A. punctipennis + A. bethuliensis + A. proboscidea.Jacknifing recovered six nodes with acceptable support in the consensus tree including 57% for the clade formed by A. angusticephala spec.nov.+ A. whocantell spec.nov., from 1000 replications.Bremer support for the major nodes on this cladogram was calculated from 2870 retained trees, with length ranging from 81 to 111 steps, and the results are as follows: Desmatomyiini, 9 steps; Desmatomyia, 14 steps; Crocidiini, 5 steps; Apatomyza, 2 steps; A. bethuliensis + A. proboscidea, 2 steps; A. whocantell spec.nov.+ A. angusticephala spec.nov., 3 steps; Mallophthiria + Megaphthiria, 2 steps; Megaphthiria, 8 steps; Crocidium, 4 steps; C. lactipenne + C. microstictum, 2 steps; C. dasypolium + C. phaenochilum, 18 steps (Fig. 10).
APPENDIX 1. Lines added (in bold) to the data matrix of 30 morphological characters used in Lamas & Couri (2005).