taxonID	type	description	language	source
03D287A0EB25E048BE48F9D4BBD3FE17.taxon	description	(Figures 1 — 6)	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB25E048BE48F9D4BBD3FE17.taxon	diagnosis	Diagnosis. Scaptomyza is a cosmopolitan genus that currently contains about 15 subgenera (Wheeler 1981, 1986), some of which have been designated as separate genera at one time or another (Frey 1954; Hackman 1959, 1982; Malloch 1934). The traditional definition of Scaptomyza includes taxa with two to four rows of acrostichal setulae, two pairs of postsutural dorsocentral setae (and sometimes with a single set of presutural dorsocentrals as well), the third costal section 2.5 times longer than the fourth, and the head distinctly longer than high (Hardy 1965). However, a rather large radiation of about 150 described species present in the Hawaiian Archipelago, has broadened this definition somewhat, mainly because of atypical characters possessed by some of these taxa (Hackman 1959, 1962, 1982). For example, many members of the subgenus Elmomyza have six rows of acrostichal setulae, suggesting that this character may be quite variable within Scaptomyza. Therefore, having either two or four rows of acrostichals is not a good synapomorphy for the genus Scaptomyza, although it may be useful at delimiting some subgenera. Perhaps the best character defining all Scaptomyza is the presence of well developed, exposed surstyli and enlarged lobes on either the epandrium (ninth tergite), cerci, or both. These morphologies are also characteristic of the genera Grimshawomyia and Titanochaeta, as well as the subgenus Engiscaptomyza. An additional character, found in females of most species, is a weakly developed, fleshy, non­dentate ovipositor. Titanochaeta is atypical in this character as females of this group have a slender, sharply pointed, stylet­like ovipositor, a character that may be an adaptation to a lifestyle as a spider egg sac predator. Methods. We have examined the types, as well as large series of other material, from all species placed in Titanochaeta, Engiscaptomyza, and Grimshawomyia (Table 1). We also have examined material from most recognized subgenera of the genus Scaptomyza. Based on this work, we selected a number of taxa placed in the genus Scaptomyza, as well as representatives of Drosophila (Engiscaptomyza), Grimshawomyia, and Titanochaeta thought to be closely related to this genus, for use in the current molecular and morphological analyses. Over 3.3 kilobase pairs of nucleotide sequence from five genes (16 S, Adh, COI, COII, Gpdh) were examined in about 120 drosophilid species using a variety of phylogenetic methods (Bonacum 2001). The phylogeny shown in figure 1 is the result of a maximum parsimony analysis (addition sequences = random, number of replicates = 100, branch swap = TBR). The search recovered four most parsimonious trees [length 33,181; CI = 0.31; RI = 0.53; see Bonacum (2001) for more detail]; figure 1 is from the strict consensus. Measures of support include bootstrap proportions (BP; Felsenstein 1985, 1988), and decay indices (DI; Bremer 1988). This phylogeny shown is part of a larger study treating phylogenetic relationships within the entire Hawaiian Drosophilidae (Bonacum 2001; Bonacum et al. in press) and includes several outgroups, as well as representatives of all major Hawaiian Drosophila lineages. Based on this taxon sampling, we feel confident in making statements concerning the relationships of the Hawaiian Drosophilidae and the genus Scaptomyza. Morphological analyses were done either using light or scanning electron microscopy. Specimens were prepared as follows: adult flies stored in 70 % ETOH were completely dehydrated via sequential washes with 80 %, 90 %, 95 % and 100 % ETOH. These specimens were then critical point dried using standard protocols (Grimaldi 1987). Male genitalic structures were dissected from the abdomen and adhered to a specimen mount (Ted Pella, Inc.) using double coated, carbon conductive tabs (Ted Pella, Inc.). The material was sputter coated and visualized using a Hitachi S 4700 Field Emission Scanning Electron Microscope. All image files were saved in. tif file format and edited in Adobe Photoshop 5.0 (Adobe Systems, Inc.). Image files are available upon request. Results and Discussion. The molecular and morphological data strongly support the notion that the genera Titanochaeta and Grimshawomyia, as well as the Drosophila subgenus Engiscaptomyza, actually belong within the genus Scaptomyza. The molecular phylogeny we present (Fig. 1) shows high support for a clade containing these three endemic Hawaiian groups with several subgenera of the genus Scaptomyza (BP = 100, DI = 24.75). Although support for the Hawaiian Scaptomyza lineage plus Titanochaeta, Grimshawomyia and the subgenus Engiscaptomyza is quite robust, relationships within this clade are not well supported. Only the monophyly of the subgenus Bunostoma (BP = 100, DI = 44.5) and the sister group relationship of S. (Scaptomyza) graminum and S. (Parascaptomyza) elmoi (BP = 100, DI = 24) are well supported (Fig. 1). The latter relationship, however, implies that the subgenus Parascaptomyza is not monophyletic. This phylogeny also calls into question the monophyly of Engiscaptomyza, placing S. crassifemur as the sistertaxon of S. chauliodon and S. nasalis as the sister of S. palata (Fig. 1). Clearly, the large Scaptomyza lineage will need to be surveyed more extensively and completely revised in order to resolve these issues .. Scanning electron microscopy was used to compare the morphology of the male genitalia of Titanochaeta, Grimshawomyia, and the crassifemur group with Scaptomyza and Drosophila. It is clear that, based on several characters, the three endemic Hawaiian groups are more closely related to Scaptomyza than they are to Drosophila. For example, the epandria and cerci of Titanochaeta, Grimshawomyia, Scaptomyza, and Engiscaptomyza are all highly modified, possessing expanded lateral lobes that often bear elongate setae (Figs. 2 – 6). Hardy (1965: 606) noticed these characters and cautioned against referring to them as secondary claspers because he preferred “ to use this term only for those distinctly clasper­like lobes ... which bear strong spines. ” In addition, the genitalia of Scaptomyza, Titanochaeta, Grimshawomyia, and Engiscaptomyza have a more “ open ” arrangement, where the surstyli and lateral lobes of the epandrium form a “ cup ” on the ventrodistal surface of the abdomen (Figs. 2 ­ 6). This is in contrast to the genus Drosophila, where the surstyli are closely oppressed on either side of the aedeagus and lateral lobes on the epandrium or anal plates are generally absent. Chromosome studies also suggest a close affinity between Scaptomyza, Engiscaptomyza and Titanochaeta (Clayton et al. 1972; Yoon et al. 1975). The metaphase configurations (1 V­shaped, 3 rods, and 1 dot; N = 5) is shared between Scaptomyza, Titanochaeta, Engiscaptomyza and some species in the modified mouthparts species group (genus Drosophila). All other Hawaiian Drosophila species have the “ ancestral ” karyotype (5 rods and 1 dot; N = 6) for the genus Drosophila. It has been suggested that this reduction in chromosome number has taken place via centric fusion events (Patterson & Stone 1952). Our molecular phylogeny (Fig. 1) suggests that this has taken place at least twice – once in the modified mouthpart species and again in the Scaptomyza lineage (which contains Titanochaeta, Engiscaptomyza, and Grimshawomyia). Based on the morphological, chromosomal, and molecular evidence, we propose placing the genera Titanochaeta and Grimshawomyia into the genus Scaptomyza as subgenera. We are also moving the subgenus Engiscaptomyza from the genus Drosophila to Scaptomyza. This placement will broaden the morphological concept of the genus Scaptomyza which will, in turn facilitate further taxonomic studies on this poorly understood and complex group.	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB2CE047BE48FEDEBF77F9A7.taxon	diagnosis	Diagnosis. Mesonotum typically with five dark brown to black vittae extending the full length, the lateral vittae may be interrupted at the suture (Kaneshiro 1969). The male genitalia of the Engiscaptomyza species are also quite similar to those of the genus Scaptomyza, characterized by large lobes of the epandria and prominent surstyli (Fig. 4). Some species (i. e., S. crassifemur) also possess enlarged, swollen femora. Included Taxa. Scaptomyza (Engiscaptomyza) ampliloba (Hardy), comb. nov. from Kaua‘i, S. (Engiscaptomyza) crassifemur (Grimshaw), comb. nov. from Maui and Moloka‘i, S. (Engiscaptomyza) inflata (Kaneshiro), comb. nov. from O‘ahu, S. (Engiscaptomyza) lonchoptera (Hardy), comb. nov. from Maui, S. (Engiscaptomyza) nasalis (Grimshaw), comb. nov. from Maui and Moloka‘i, S. (Engiscaptomyza) reducta (Hardy), comb. nov. from Hawai'i, S. (Engiscaptomyza) undulata Grimshaw comb. nov. from Hawai‘i (Nishida 2002)	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB2CE047BE48FEDEBF77F9A7.taxon	discussion	Discussion. Kaneshiro (1969) recognized a single species group (crassifemur) containing two subgroups (crassifemur and nasalis) within this subgenus. We will leave all taxonomic ranks below the level of subgenus intact. The polytene chromosome of the subgenus Engiscaptomyza, unlike those of the remainder of the genus Scaptomyza, are large and easy to decipher. Yoon et al. (1975) examined the phylogenetic relationships between four species of the subgenus Engiscaptomyza using polytene chromosome banding patterns. Their work suggests that S. reducta (Hawai‘i) and S. crassifemur (Maui Nui) are sister taxa. Scaptomyza inflata (O‘ahu) is basal to this group and S. ampliloba (Kaua‘i) is the most basal member of this subgenus. KEY TO SPECIES OF THE SUBGENUS Engiscaptomyza Kaneshiro	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB2DE045BE48F956BF77FE6F.taxon	diagnosis	Diagnosis. The genus Grimshawomyia was described by Hardy (1965) and contains two species, G. palata and G. perkinsi, the latter of which was initially described as a member of the genus Drosophila by Grimshaw (1901). The unique male genitalia, which feature surstyli that are exposed and an epandrium which is developed into a pair of moderately large lateral lobes, extending beyond the apices of the surstyli (Figs. 5 ­ 6), suggest that these taxa actually form a clade within the genus Scaptomyza. This clade is also characterized by having the second antennal segment sharply pointed at the apex, extending over the base of the third segment; the vertical and ocellar setae inserted into the somewhat swollen sides of the vertex; a long costal fringe, which extends nearly to vein R 4 + 5, and the distinctive wing markings. Included Taxa. Scaptomyza (Grimshawomyia) palata (Hardy), comb. nov. from Maui and O‘ahu, and S. (Grimshawomyia) perkinsi (Grimshaw), comb. nov. from Maui, O‘ahu and Hawai‘i (Nishida, 2002). KEY TO SPECIES OF THE SUBGENUS Grimshawomyia Hardy	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB2FE043BE48FD9EBF77FC4F.taxon	diagnosis	Diagnosis. Titanochaeta was erected by Knab (1914) as a genus of endemic Hawaiian Drosophilidae. Hardy (1965) suggested that this genus actually should be considered a subgenus of Scaptomyza based on a variety of morphological characters including conspicuous surstyli which project well beyond the margins of the ninth tergite (Fig. 3), lack of ventral rays on the arista, and a short head which is approximately two times higher than long. We are placing the eleven known species of Titanochaeta, all of which are parasitic on spider egg sacs, in a subgenus of Scaptomyza. Chaetotaxy is an important synapomorphy of the subgenus Titanochaeta. These species have eight rows of acrostichal setulae, a character not seen in the other species of Scaptomyza. Furthermore, the setae on the head and thorax are very strong, the vertical setae are often longer than the head is wide. The shape of the head is also characteristic. It is equal to or narrower than the thorax in width, with a distinctly oblique, slanted front which is often more than two times longer than the lower margin of the head. The genae are straight sided and indented along the anterior eye margin. Finally, the ovipositor of most taxa is long and needle­like, probably due to the parasitic lifestyle of these taxa.	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
03D287A0EB2FE043BE48FD9EBF77FC4F.taxon	discussion	Discussion. The new subgeneric status of Titanochaeta in Scaptomyza requires that we propose new replacement names for T. evexa, T. kauaiensis, and T. silvicola because those specific epithets are preoccupied in the genus Scaptomyza. We propose: Scaptomyza neoevexa O’Grady et al., new replacement name for Titanochaeta evexa (Hardy, 1965) (preoccupied by Scaptomyza evexa Hardy, 1965), Scaptomyza neokauaiensis O’Grady et al., new replacement name, for Titanochaeta kauaiensis (Hardy, 1965) (preoccupied by Scaptomyza kauaiensis Hackman, 1959), and Scaptomyza neosilvicola O’Grady et al., new replacement name, for Titanochaeta silvicola (Hardy, 1965) (preoccupied by Scaptomyza silvicola Hardy, 1965). Included Taxa. Scaptomyza (Titanochaeta) bryani (Wirth), comb. nov. from Hawai‘i, O‘ahu, and Maui Nui, S. (Titanochaeta) chauliodon (Hardy), comb. nov. from Maui and O‘ahu, S. (Titanochaeta) contestata (Hardy), comb. nov. from O‘ahu, S. (Titanochaeta) glauca (Hardy), comb. nov. from Maui, S. (Titanochaeta) ichneumon (Knab), comb. nov. from Hawai‘i, S. (Titanochaeta) neoevexa O’Grady et al., from Moloka‘i, S. (Titanochaeta) neokauaiensis O’Grady et al., from Kaua‘i, S. (Titanochaeta) setosiscutellum (Hardy), comb. nov. from Hawai‘i and Moloka‘i, S. (Titanochaeta) neosilvicola O’Grady et al., from Hawai‘i, S. (Titanochaeta) sweyzei (Wirth), comb. nov. from Kaua‘i, Maui, and O‘ahu, and S. (Titanochaeta) vittigera (Hardy), comb. nov. from Kaua‘i (Nishida, 2002). Discussion. The species in this group are infrequently collected and poorly understood taxonomically. The fact that a number of species in the subgenus Titanochaeta are present on multiple, non­adjacent islands suggests that they are either better at dispersing or have a lower rate of speciation than other endemic Hawaiian groups. It also might indicate that there are additional cryptic species remaining to be described in this group. KEY TO SPECIES OF THE SUBGENUS Titanochaeta Knab	en	O’Grady, Patrick, Bonacum, James, Desalle, R., Val, Francisca Do (2003): The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159 (1): 1-16, DOI: 10.11646/zootaxa.159.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.159.1.1
