Table Of Content)
PROC. ENTOMOL.SOC.WASH.
1()5(1),2003,pp. 30-49
A REVISION AND PHYLOGENETIC STUDY OF LIPOCHAETA COQUILLETT
(DIPTERA: EPHYDRIDAE)
Wayne N. M.athls and Michelle D. Trautwein
(WNM) Department of Systematic Biology. Entomology Section, P.O. Box 37012,
NMNH. MRC-169. Smithsonian Institution.Washington.DC20013-7012.U.S.A. (e-mail:
mathis.wayne@'nmnh.si.edu): (MT) 2845 Shoal Crest. Auistin.TX 78705.U.S.A. (e-mail:
[email protected]
—
Ahstnict. The genus Lipochacia Coquillett is revised and a phylogenetic analysis of
the genera ofthe tribe Lipochaetini is provided. Lipochaeta is known thus faronly from
the New World, where there are now two species. The second species. L. ranica, n. sp..
(type locality: California), is described herein. The species ofLipochaeta occur in saline
or alkaline habitats, especially along maritime coasts. The monophyly of Lipochaeta is
well corroborated and its putative sister group is the Old World genus HomaloDietopluis
Becker
Kex Words: revision. Diptera. Ephydridae. Lipochaetini. L. ranica. New World, phylog-
eny
Among 114 genera of shore flies (Dip- provide a subfamilial name. Becker (1896:
tera: Ephydridae), less than 17% (19 gen- 275) quoted Coquillett"s entire paper but
era)aremonotypic(MathisandZatwarnicki substituted "Lipochaetinae" forLipochaeta
1995). and that percentage is decreasing as in the introductoryportion,thus makingthe
we better sample the extant fauna and dis- subfamilial name available. Attribution of
coveradditional species. Field work during Lipochetinae as a family-group name to
the last two decades, for example, has re- Becker was followed in all recent catalogs
vealed hundreds of new species but very and is continued here. Sabrosky (1999:
few new genera. Thereisalsoadecreasein 179), however, suggested that. "It seems
monotypic generaas we clarify and rechar- reasonable to correct the obvious lapse and
acterize higher level taxato be more inclu- credit Coquillett with the subfamily." Co-
sive, monophyleticclades. usuallycompris- quillett clearly was the first person to rec-
ing more than one species. The genus Li- ognize these taxa. from species to subfam-
pochaeta Coquillett. the subject ofthis re- ily, but the subfamilial nameiscorrectlyat-
vision, is an example of this evolving tributed to Beckerand the generic and spe-
pattern in the classification ofshore flies. cies names to Coquillett.
Coquillett (1896) described Lipochaeta The bizarre external appearance of Li-
slossonaeinthelate 19thCentury,anduntil pochaeta, being highly adapted to psam-
now it was the only included species in the mophilous habitats, initially confused some
genus. Coquillett recognized that this new authors about its familial affiliation. A year
genus and species were unusual and appro- afteritsdescription.Williston(1897: 8)pre-
priately suggested that they be placed in a ferredplacementofLipochaeta ". . .among
separate and new subfamily, but he did not the Ochthiphilinae in the vicinity of Rhic-
VOLUME 105.NUMBER I 31
itoessa [Tethinidae]" and added that. "No to Lsganiera Giordani Soika andAsmeringa
Ephydrid that I know of lacks bristles, Becker,twoOld-Worldgenerathatoccuron
while both ofthese latter families [Oscini- theseashoresofthe MeditenaneanandEast
dae and Agromyzidae] have numerous Africa. Giordani Soika further suggested
forms without them. The face is too short, that the occurrence of Lipochaetini in the
theantennaetoodifferentinstructuretobe- New Worldwasduetocontinental driftand
long with the Ephydridae. Moreover the that the group has greater antiquity than
pollinose body and white wings, while not was previously thought. Mathis (1984a)
absent among Ephydridae, are not at all concurred with Giordani Soika inremoving
common."" Townsend (1898: 168) de- Lipochaeta from Parydrinae and in placing
scribed a second species in Lipochaeta (L. it close to Lsgamera and Asiiwringa in the
texeusis from Padre Island, Texas) and subfamily Gymnomyzinae.
commented on the ta.xonomic placement of The above-cited studies were based on
the genus. Townsend wrote that while Li- phenetic or overall differences and similar-
pochaeta is "". . . clearly alliedto the Ephy- ities, and although the placement of Lipo-
dridae" it ". . . is truly one ofsingular as- chaeta did change from Parydrinae to
pect and anomalous position."" Aldrich's Gymnomyzinae. its position close to Isga-
(1905)catalogofNearctic Dipterafollowed niera and Asmeringa is unsupported. Al-
Coquillett and Townsend in listing Lipo- though the latter two genera appear similar
cliaeta as a genus in the Ephydridae and externally, these features represent, for the
also in placing L. texensis as ajunior syn- most part, independent and convergent ad-
onym ofL. slossonae. The synonymy ofL. aptations to psammophilous environments
texensis with L. slossonae was apparently that are associated with seashores, not syn-
based on information in a letterthat Willis- apomorphies that indicate phylogenetic re-
ton had written to Aldrich. Although Jones lationships.
(1906)did notacceptLipocluieta asashore Another advance in the phylogenetic po-
fly in his worldwide catalog ofEphydridae. sition and composition ofLipochaetini was
he offered no alternative placement. Nearly the recognition that the tribe Atissini. as
all subsequent authors have followed Co- thencharacterized,waspolyphyleticandin-
quillett. Townsend. and Beckerin recogniz- cluded genera that are more closely related
ing Lipochaeta as an ephydrid. usually in to Hecainede Haliday (tribe Hecamedini,
the tribe Lipochaetini. subfamily Parydri- Mathis 1993) and to Lipochaeta (tribe Li-
nae (Stuilevant and Wheeler 1954; Wirth pochaetini. Zatwarnicki 1992. Mathis
1965. 1968: Mathis 1977: Cogan 1980). A 1995a). Zatwarnicki (1992) provided evi-
notableexception wasE.T Cresson,Jr.the denceindicatingthatthetribe Atissini isre-
doyen of 20th Century shore-fly workers, lated to taxa in the subfamily Hydrelliinae.
who was silent on the subject. whereas Hecamedini and Lipochaetini are
Although recognition of Lipocluieta as in the subfamily Gymnomyzinae. Zatwar-
an ephydrid is now virtually universal, its nicki (1992) cited furtherevidence that He-
placement in available shore-fly classifica- camedini and Lipochaetini are sister
tions, especially in catalogs, has varied. groups. The principal source of characters
Cresson's successors (Sturtevant and (synapomorphies) for these studies derives
Wheeler 1954: Wirth 1965. 1968: Mathis from structures ofthe male terminalia. The
1977: Cogan 1980)accordedtribal statusto latter studies, especially Zatwarnicki
the genus in the subfamily Parydrinae. ap- (1992). indicate that the tribe Lipochaetini
parently considering Lipocluieta to be re- is closely related to the tribe Hecamedini
lated to the tribes Parydrini and Hyadinini. and includes the following four genera
Giordani Soika(1981). however, wasofthe (date and author(s) who first placed the ge-
opinion that the tribe Lipochaetini isrelated nus in Lipochaetini are noted in parenthe-
—
PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
sis): Glenanthe (Mathis and Zatwarnicki as part of generic treatments and phyloge-
1990b), Paragleiuifithe Wirth (Zatwarnicki neticconsiderationstodocumentthemono-
1992). Lipochaeta (Becker 1896). and phyly ofthe lineages, particularly at thege-
Homalometopiis Becker (Mathis 1984b). neric level.
Recent revisionary and phylogenetic stud- Although many specimens for this study
ies of genera of the tribe Lipochaetini in- are in the National MuseumofNatural His-
clude Homalometopiis (Mathis 1984b, Mu- tory. Smithsonian Institution, Washington,
nari 1988) and Glenanlhe (Mathis 1992). DC (USNM),wealsoborrowedandstudied
numerous specimens that are deposited in
Methods and Materials
the following museums:
Thedescriptiveterminology,withtheex- ANSP Academy of Natural Sciences of
ceptions noted in Mathis(1986)and Mathis Philadelphia, Pennsylvania, USA.
and Zatwarnicki (1990a), follows that pub- AMNH American MuseumofNatural His-
lished in the Manual ofNearctic Diptera tory, New York. USA.
(McAlpine 1981). Because mspmecimens are BMNH The Natural History Museum (for-
small, usually less than 3.5 in length, merly the BritishMuseum(Natural
study and illustrationofthe maleterminalia
History)). Li>ndon. England, Unit-
rWeequhiraevdefuosleloowfedathceomtepromuinndolomgiycrfoosrcmoopset. CNC eCdanKaidinagndoNma.tional Collection. Ot-
structures ofthe male terminalia that other
tawa. Canada.
workers in Ephydridae have used (see ref-
erences in Mathis 1986; Mathis and Za- Systematics
twarnicki 1990a. b). The terminology for Tribe Lipochaetini Becker
structuresofthemaleterminaliaisprovided
directly on Figs. 17-18. The species de- Lipochaetini Becker 1896: 275 [as Lipo-
scriptionsarecompositeandnotbasedsole- chaetinae]. Type genus: Lipochaeta Co-
ly on theholotypes. One headandtwoven- quillett 1896.—Zatwarnicki 1992: 89,
ational ratios that are used in the descrip- I18-119 [listingofinc—ludedgenera,phy-
tions aredefined below (all ratios arebased logenetic placement]. Mathis and Za-
on three specimens: the largest, smallest, twarn—icki 1995: 160-163 [world cata-
and one other). Gena-to-eye ratio is the ge- log]. Mathis 1995a: 2-4 [description,
nal height measuredatthema.ximumheight key to genera].
divided by the eye height. Costal vein ratio Diagnosis (synapomorphies indicated by
isthe straight linedistancebetweentheapi- an asterisk (*)). Head: Frontal vitta (or
cesofR2+,andRj.jdividedby thedistance ocellar triangle) setulose*; ocellar seta ei-
M
between the apices ofR, and R:+v vein ther greatly reduced or absent (sometimes
ratio is the straight line distance along vein with a pair of intrafrontal setae slightly
M
between crossveins (dm-cu and r-m) di- larger than other setulae. this pair inserted
vided by the distance apicad ofdm-cu. in front ofanterior ocellus)*: pseudoposto-
Thephylogeneticanalysiswasperformed cellar seta reduced or lacking*: fronto-or-
with the assistance of Hennig86iC>, a com- bital setae 3 (reduced secondarily in Lipo-
puterized algorithm that produces dado- chaeta). anterior 2 setae proclinate. poste-
grams by parsimony. Character data were rior seta reclinate. Pedicel lacking promi-
polarized primarily using outgroup proce- nent, spinelike seta; arista with cuticular
dures. Although autapomorphies were not hair dorsally and ventrally. appearing ma-
included in thecladistic analysis(they were cropubescent or brushlike, without dorsal
made inactive), which would skewthecon- rays*. Eye bearing numerous interfacetal
sistency and retention indices, we listed microsetulae (apparently arising from each
them on the cladosram and included them interfacet). Genal seta reduced or lacking.
VOLUME 103. NUMBER 1
Tliorax: Dorsocentral setae weakly de- or polarity of the few characters that may
veloped, only posteriormost pair conspicu- indicate relationships. For example, a gap-
ous; acrostichal setulae in 2-4 rows, fre- ingoral cavity andawideclypeusarechar-
quently with a prescutellar pair better de- acter states that are common toLipochaeta
veloped: postsuturalsupra-alarsetalacking: andHomalometopus, whereasanarroworal
frequently postpronotal and presutural opening and clypeus occur in Glenaiithc
supra-alarsetareducedorlacking;posterior and Paraglenanthe. The problem is that
notopleural seta inserted above level ofan- both the narrow and wide conditions occur
terior seta, sometimes only slightly so (as in the outgroup, Hecamedini, as well as
in Homalometopus and Lipoduieta). Wing commonly elsewhere in the Ephydridae,
with vein ^2 long,extended nearly to lev- thusconfusing issuesaboutwhichcharacter
7.
el of apex of vein R4+5. Legs lacking con- stateisapomoiphicwithin the Lipochaetini.
spicuous setae; femora and tibiae usually The tribe Lipochaetini. wiiich is one of
gray to pale brown; tarsi yellow. six tribes now placed in the subfamily
Ahdonieu: 5thtergiteofmalelongerthan Gymnomyzinae (Mathis and Zatwarnicki
4th. Male terminalia as follows: epandrium iyy3), appears to be mostclosely related to
attenuate, either emarginate posteriorly or the tribe Hecamedini (Zatwarnicki 1992).
incomplete dorsally; surstylus well devel- Lipochaetini'ssister-grouprelationshipwith
oped, usually elongate, frequently as long Hecamedini is corroborated by two syna-
or longer than epandrium; aedeagus elon- pomoiphies that we have identified (includ-
gate, slender, tubular, apex with recurved ingZatwarnicki's [1992|characters59-60):
flap oriented posterodorsally, apical flap in 1) pre- and postgonites apparently fused or
groove at rest, base of aedeagus bifurcate, greatly reduced; 2) posterior notopleural
sometimeswitharmselongate*;ejaculatory seta inserted much fartherdorsad from no-
apodemepresent,compressedlaterally*;ae- topleural suture than anterior seta.
deagal apodeme L-shaped, sometimes with Lipochaetini are distinguished from He-
extended ventromedial process; gonites camedini and other tribes of the subfamily
(pre- and postgonites) lacking, possibly Gymnomyzinae and the tribe's monophyly
fused with hypandrium*; hypandrium well is confirmed by the following characters
sclerotized, usually V- or U-shaped: 3th (synapomorphies are noted by an asterisk
sternite deeply V- or U-shaped into which (*)): 1) body densely invested with micro-
the surstyli and ae—deagus lie at rest. tomentum, generally dull colored; *2)ocel-
Natural history. This tribe is unusually lar seta lacking or weakly developed; *3)
tolerant of alkaline or saline aquatic envi- pseudopostocellar seta lacking; *4) arista
ronments, and speciesofmostgeneraoccur microsetulose,lackingdorsalrays;*5)fron-
on seashores or are associated with inland tal vitta bearing many setulae; 6) posterior
aquatic habitat—s that are saline or alkaline. margin of gena broadly rounded onto oc-
Discussion. Although the tribe Lipo- ciput; 7) scutellum with 2 pairs ofmarginal
chaetini andeachoftheincludedgeneraare setae; 8) foreleg normal, forefemurslender,
readily characterized, often with substantial foretibia not having a large spur apically;
evidencethattheyaremonophyletic,there- 9) abdominal tergites 2-4 subequal in
lationships among the genera are not well width, microtomentose, but more or less
understood. In the key that follows, forex- smooth; *10) aedeagus with apical flap or
ample, we have included numerous char- appendix that is folded back dorsally: and
acters, nearly all autapomorphies, that fa- *11) gonite reduced.
cilitate identification ofthe genera but con- With the phylogenetic background ofthe
tribute virtually nothing to resolution of tribe Lipochaetini within the subfamily
their phylogenetic relationships. A further Gymnomyzinae established and the mono-
complicating factor is the debatable status phyly ofLipochaetini documented, we now
n
34 PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
—
Hecamede
1 9 10 16
110-D— Glenanthe
2 4 17
1 1 1
Paraglenanthe
10 16 19 20
1111
5 6 11 12 14 15 17 18 19 20
12 12 10-D-a Lipochaeta
7 8 14 1 2
1 1 1
Homalometopits
Fig. I. Cladogram depicting hypothelical clailistic lelalinnships among genera ofLipochaetini (length 17
steps,consistencyindex0.82; retentionindex0.62).
proceed with the cladistic analy.sis and re-
sultant relationships among the included
genera, but with afew explanatoryremarks
first. In the presentation on genus-level re-
lationships that follows, thecharactersused
in the analysis are noted first. Each char-
acter is immediately followed by a discus-
sion toexplain its statesandtoprovideper-
spective and any qualifying comments
about that character After presentation of
theinformationoncharacterevidence,ahy-
pothesisofthecladistic relationshipsispre-
sented and briefly discussed. The clado-
gram (Fig. 1) is the primary mode to con-
vey relationships, and the discussion is to
supplement the cladogram and is intended
only to complement the latter. In the dis-
cussion of character data, a "O"" indicates
Table 1. Matrixofcharactersandtaxausedinthe
cladistic analysis ofLipochaetini (numbers for char-
acterscorrespondwiththoseusedinthetext).
)
VOLUME 10?.NUMBER 1 35
4(4). Oral opening: (0) wide, gaping; (1) for Honialometopns and Lipochae-
narrow (a synapomorphy for Glen- ta).
cmthe and Paragleiuiiithe). 5(14). Setae along posteriormarginofane-
3(5). Height of face: (0) normal, usually pisternum: (0) 2 setae and some
higher than wide; (1) short (an au- smaller setulae; (1)1 seta (an auta-
tapomoiphy for Lipochaeta). pomorphy forHonialoinetonuis); (2)
6(6). Size of antenna: (0) normal; (I) re- no large setae, only setulae (an au-
duced, especially the arista (an au- toapomorphy for Lipochaeta).
tapomorphy forLipoclniela). 6(15). Wing near apex of subcostal vein
1{1). Distance between antennal bases: and vein R,: (0) normal, membra-
(0) narrow, distance between anten- nous; (1) sclerotized and thickened,
nal bases less than antennal width; yellow (an autapomorphy for Lipo-
(1) wide, antennal bases separated chaeta).
by widthgreaterthanantennalwidth 7(16). Wing membrane coloration: (0) hy-
(an autapomorphy for Hoinalome- aline, transparent; (I) white, trans-
topits): (2)verywide,antennalbases lucent (a synapomorphy for Glen-
separated by 3X antennal width (an anthe. Paraglenanthe. and Lipo-
autapomorphy forLipochaeta). chaeta).
8(9). Shape ofmesofrons: (0) narrow. V- Abdomen
shaped. with wide parafrons and
fronto-orbits (Glenanthe and Para- 1(17). Epandrium: (0) entire dorsally; d)
glenanthe): (I) wide, broadly U- attenuate with a posterodorsal notch
shaped. parafrons and fronto-orbits (an autapomorphy for Glenanthe
comparatively smaller (a synapo- and Paraglenanthe): (2) incomplete
morphy for HoDialonic'topus and Li- dorsally withagapbetween twolat-
pochaeta). eral portions (an autapomorphy for
9(9). Height of gena: (0) high, 0.33 to Lipochaeta).
more than 0.5X eye height; (I) 2(18). Cerci: (0) separate, unfused with
short, less than Vg eye height (an au- medial margin of epandrium: (I
tapomorphy for Glenanthe). fused laterally with median margin
ofepandrium (an autapomorphy for
Thorax Lipochaeta).
3(19). Length of surstylus: (0) about same
1(10). Katepisternal seta: (0) present, con- length as epandrial length; (1) much
spicuous, well developed; (I) great- longerthan epandrial length,usually
ly reduced (a synapomorphy for by more than twice (a synapomor-
Homalometopiis. Lipochaeta. and phy forHonialometopns. Glencmthe.
Paraglenanthe). and Paraglenanthe).
2(11). Postpronotalseta: (0)present: (I)re- 4(20). Base of aedeagus: (0) truncate or
duced (an autapomorphy for Lipo- nearly so; (1) bifurcate (a synapo-
chaeta). morphy for Honialometopns and
3(12). Presutural supra-alar seta: (0) pre- Paraglenanthe); (2) arms of basal
sent; (I) absent (an autapomorphy bifurcationmoreelongateandcurled
for Lipochaeta). beneath (an autapomorphy for Li-
4(13). Position of posterior notopleural pochaeta).
seta; (0) inserted above level ofan-
terior seta; (1) inserted at about Analysis and Results
same level as anterior seta or t)nly Using the implicit enumeration (ie*) op-
slightly elevated (a synapomorphy tion of Hennig86, which is an exhaustive
PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
Table 2. Analysis ofcharacters based on the cladogram (Fig. I). Con. Index ^ Consistency Index: Rel.
Index = RetentionIndex.
—— —
VOLUME 105. NUMBER I
—
a good indicator. Panigleiuintlie. on the 1905: 631 [Nearctic catalog]—, Jones
otherhand, ha.s a more limiteddistribution, 1906: 169 [note, diagnosis). Curran
occuiTing only in the New World where it 1934:—346-347 [tigs, of head, generic
is primarily circumcaribbean. key]. Sturtevant and Wheeler 1954:
215-216 [listedas agenus in Napeae].
Ke'> to Genera of Lipoch.aetini Becker Wirth 1965: 750 [Nearctic c—atalog];
1. Eye pyriform, distinctly narrowed ventrally; 1968: 22 [Neotropical catalog]. IVIathis
genashort,lessthanone-fourtheyeheight;ka- and Zatwarnicki 1995: 163 [world cata-
tepisternalsetapresentalongposteriormargin, log].
moderately welldeveloped . . GleiuinlhcHaliduy
- Eye[15genseprecailelsy,owvoarllodrwirdoeu;ndM,antohtisdis1t9i9n5catl|ynar- Diagnosis, Lipochaeta is distinguished
rowed ventrally; genahigh,one-thirdormore from other genera of Lipochaetini by the
ofeyeheight;katepisternalsetareduced .... 2 following combination ofcharacters: Body
2. Antenna reduced, inserted in well-separated setae and setulae generally inconspicuous,
cavity, arista atrophied, budlike; face short, pale. Antennae widely separate and arista
height subequal to lengthofreduced antenna; rudiinentary. Frons projected and large;
cseltyapeeuasndbasentdulliakee;paloer;alcoostpaendiinstgincgtalpyintgh;ickbeondeyd sub-cranial cavity large, gaping. Body
atmergerofveinR, Li[>c>cluiciaCoquilletl length 2.0-4.0 mm. generally grayish, dor-
|2species.NewWorld;mostlycostalmarine sum ofhead and thorax sometimes brown-
onsandbutalsoinlandwheresalineconditions ish, and almost entirely microtomentose.
- eAxnitste)nnanormallydeveloped,not indeepcav- Description. Head: Wider than high in
diteyv,ealroi,psetda,asheliognhgtamsufclhagemlolroemertehan1; lleancgethwelolf amnetnetroiosre,visepwa,rsgerlayyisshe;tuleonstier;elymesmiocfrrootnos-
antenna;clypeusvariable;oralopeningnarrow wide, broadly U-shaped, uniformly setulo-
orgaping;setaeandsetulaelargelyblack;costa se, differentiated from remainder of frons
3. Monelsyofsrloignhstlylatrhgiec,keonecdcuaptymienrggemrosoftveoifnIRr,ons.,. 3 by darker gray to brownish gray; parafrons
platelike,subrectangular,uniformlyandevenly a comparatively small triangular anterior
setulose; ventral facial margin flat; clypeus area; fronto-orbits comparatively narrow.
wide,bandlike;oralopeninglarge,gaping . . Head without conspicuous setae. Ocellar
HoinalimiclD/nisBecker setae absent; pseudopostocellar setae ab-
Mat[h7isspe1c9i8e4sb..EMaustnearrniH1e9m8i8s)phere (Meditenanean); sent, fronto-orbital setae absent (reduced
- Fronslackingdifferentiatedmesotrons,atmost secondarily); ocelli arranged to form equi-
with frontal orocellartriangle or vitta that is lateral triangle. Antenna reduced, in well-
weaklydifferentiatedfromremainderoffrons; separated (gap between antennal bases
ventral facial marginemarginate;clypeusnar- about 3x antennal width), deep cavities,
row,exposedthroughventral facialemargina-
tion[;3osrpaelcioepse.niNnegwsmWaolrlld.(.CaPraiibabgetaenn)c;inWtihrelhWirth tourliaeentoendlvaetnetrraallly;anpdedidcoerlsawlitshidefsi,ne,buptalleacske--
I950>1 ing dorsoapical seta, aristarudimentaryand
budlike. Eye generally irregularly round
Genus Lipocliuehi Coc|uillett with distinct, angularmarginposterodorsal-
Lipoclnicrci Coquillett 1896: 220. Type spe- ly near vertex; interfacetal microsetulae
cies: Lipochaeta slossonae C—oquillett present,numerous,conspicuous. Faceshort,
1896. original designation. Becker height subequal to length ofreduced anten-
1896: 274—-275 [quote of original de- na, narrowly triangularin profile,taperedto
scription]. Williston 1897: 7 [placement acute angle posteroventrally; clypeus band-
near Rhicnoessa, family Agromyzidae, like, wide. Gena bearing fine, pale setulae.
subfamily "Ocht—hiphilinae"]: 1908: 306 Subcranial cavity large and gaping.
[generic key]. Townsend 1—898: 168 Thorax: Entirely grayish to brownish,
[notes on generic placementj. Aldrich microtomentose. pleural areagenerally pal-
PROCEEDINGSOFTHEENTOMOLOGICALSOCIETYOFWASHINGTON
er than dorsal coloration: mesonotum bear- less Y-shaped in ventral view, with forked
ing numerous, fine, pale setulae. setulae portion basal, lateral phalanges from basal
most conspicuous on posterior portion of arms offork. Female Terminalia: Segments
scutellum. particularly in females: post- 6-7 telescoped out to form tube: 8th seg-
pronotal seta reduced: posteriornotopleural ment apparently lacking: hypoproct Y-
seta at only slightly elevated position rela- shaped. —
tive to anterior seta: anepistemum bearing Remarks. The poorly developed epan-
numerous setulae: katepistemal seta vari- drium and an aedeagus with a terminal
able. Wing with costal vein extended to membranous flap are similar to Glenanthe
veinM:costadistinctlythickenedatmerger and Homalometopiis Becker. The female
ofvein R,: dorsal setulae along costal vein terminalia is also similar to that ofHoma-
ended just beyond vein R^^,: wing white, lometopiis.
translucent, wing venation and halter yel- Until now, Lipochaeta has been mono-
low. Midfemurbearing comblike row of6— typic, with L. slossonae as the only includ-
7 white, longersetaeanteriorly atapical Vs- ed species. The second species. L. ranica,
Vi\ tarsi yellowish: midtarsus with basitar- is very similar and obviously closely relat-
somere longer than remaining four: each ed. Differences between these two species
tarsomere with dark, ventroapical setae: are seemingly slight but are consistent and
midtarsus bearing twice as many black se- significant(seekeyanddescriptionsofspe-
tae as others, these paired at apices of tar- cies below). In additiontodescribingasec-
someres. basitarsomere bearing 4-5 pairs, ond species, which is found primarily on
apical tarsomere variable: pulvilluspresent. thewestcoastofCaliforniaandMexico,we
Abdomen: Male: tergites 2-4 about also report the possibility ofa third species
equalinlength: lengthof5thtergiteslightly from Chile. The potential third species is
more than combined length of3rd and 4th represented by two female specimens from
tergites: 5th tergite triangular in dorsal Atacama. One of the females has an elon-
view: sternites 2-4 linear, much narrower gate scutellum and bears a fringe of long,
than long, weakly sclerotized: 5th stemite whitesetulaealongthemargin.Anelongate
deeply U-shaped and better sclerotized scutellum also occurs in L. slossonae and
along inner margin ofU. Male Terminalia: L. ranica, and to a degree, the scutellar
epandrium greatly reduced, in lateral view fringe ofsetulae is also apparentwithin the
about as long as cerci. fused dorsolaterally variation ofthese two species. Thus we are
with cerci. incomplete dorsally. with a gap unsureaboutthe statusoftheChilean spec-
between lateral portions: surstylus digiti- imens, i.e.. is the variation in the scutellar
form. moderately to conspicuously elon- features intra- or interspecific. Additional
gate, mostlyparallel-sided,usuallyshallow- specimens, especially males, are needed to
ly curved in lateral view, generally evenly assess betterthese possibilities.
setulose. setulae longerapically:gonitesei-
ther indistinguishably fused with h\pan- Key TO Species ofLipochaeta
drium or lacking: ejaculatory apodeme ev- 1. Tarsi often entirelyyellow; surstyluselongate
ident as a simple, slightly angulate, long, (distinctly longer than height ofepandrium).
narrow process: aedeagus very long, nar- taperedapicallytonarrowhroundedapex(east
coast of North .America from Massachusetts
row, tubular, length twice orslightly longer southtoFloridaandTexas,islandsoftheCa-
than surstylus, apex with membranous flap ribbean. Belize [Stann Creek], Mexico [Chia-
folded back on itself, length of flap about pas], and Panama [PlayaSantaClara] ....
V4-V6 length of aedeagus. basal portion of L. slossonaeCoquillett
aedeagus angled ventrally and forked with - Apical tarsomere always slightly to distinctly
darkened;surstylusshorter(subequaltoheight
aedeagal apodeme between arms of basal ofepandrium).parallelsidedtobluntlyround-
fork: hypandrium uell sclerotized. more or ed apex (west coast of North America from
—
VOLUME 105.NUMBER 1
Figs. 2—7. Scanning electron micrographs ofLipochaeta slossonae. 2, Head, anterodorsal view. 3. Same,
lateralview.4.Frons,anterodorsalview.5.Face,anteriorview.6,Rightantenna,anteriorview.7.Leftcompound
eyeand interfacetalsetae,anteriorview. Scalebarsequals 100jjim.
San Francisco south to the Mexican states of Florida, Maryland, New Jersey, Texas,
BajaCaliforniaSurandNayarit.andtotheGa- comments on habitat and behavior].
lapagosIslands) L. ranica. new species
Wirth 1956: 18 [list. Bahamas]; 1965:
Lipochaeta slossonae Coquillett 751 [Nearcticcatalog (partim)]; 1—968: 22
(Figs. 2-19) [Neotropical catalog (partim)]. Foote
Lipochaeta slossonae Coquillett 1896: 1995: 422 [sandy beach habitat].—Math-
220.—Becker 1896: 275 [description|.— is and Zatwarnicki 1995: 163 [worldcat-
Aldric—h 1905: 631 [Nearcticcatalog(par- alog (partim)].—Mathis 1997: 28-29 [re-
tim)|. Jones 1906: 169 [note, diagno- view, Belize].
sis).—Johnson 1913: 86 [list Florida].— Lipochaeta texensis Townsend 1898:
Sturtevant and Wheeler 1954: 216 [list. 168.—Aldrich 1905: 631 [synonymy].—
