New species of leptohyphidae (Ephemeroptera) from Costa Rica

2007·5.3 MB·

by  David E Baumgardner

PROC. ENTOMOL. SOC. WASH.
109(2), 2007, pp. 416^26
NEW SPECIES OF LEPTOHYPHIDAE (EPHEMEROPTERA) FROM
COSTA RICA
David E. Baumgardner
A&M
Department of Entomology, Texas University, College Station,
TX 77843-2475, U.S.A. (e-mail: [email protected])
—
Abstract. Three new species of leptohyphid mayflies (Ephemeroptera: Leptohy-
phidae), Leptohyphes mandibiilus, Tricoiythodes kirki, and Tricorythodesprimus, are
described from Costa Rica based upon the larval stages. Leptohyphes mandibuhies
can be distinguished from other species in the genus by its unique mandibular
morphology and color patterns of the operaulate gill. Tricorythodes kirk larvae are
distinctive due to their contrasting abdominal coloration. Tricorythodes primus is
distinguished from other speices of Tricorythodes in Central America due to the
presence of a distinctive and well developed frontal shelf and genal projections.
Key Words: Ephemeroptera, Leptohyphidae, Leptohyphes, Tricorythodes, new
species
The mayfly family Leptohyphidae is Materials and Methods
a diverse assemblage of more than 125 Larvae were collected from streams
species endemic to the Western Hemi- using forceps and by disturbing the
sphere. The family is most diverse in sediment using kick nets, then pre-
South America, with fewer species served in 70% ethyl alcohol. Legs,
known from Central and North Amer- gills, and mouthparts were removed
ica. The family has received critical and mounted on slides for detailed
attention recently in both South Amer- study. Figures were drawn using
ica (Molineri 1999, 2001a,b, 2002, a camera lucida. Collection sites are
2003a,b, 2004; Molineri and Zufiiga given in longitude/latitude coordi-
2004), and North and Central America nates as degrees, minutes, seconds,
(Baumgardner and McCafferty 2000; and were determined using a hand-
Wiersema and McCafferty 2000, 2003, held global positioning system unit.
2005; Wiersema et al. 2001; Baumgard- Setal descriptive terminology follows
ner 2003; Baumgardner et al. 2003, Baumgardner and Avila (2006). Gill
2006; Baumgardner and Avila 2006). formula follows Molineri (2003b), and
Additional descriptions and revisions of indicates the number of membranous
species are still required before generic lamellae on abdominal segments two
boundaries and relationships can be through seven. Collections (and their
fully analyzed. Examination of collec- acronyms) housing materials used in
A&M
tions of mayflies made in 2001 from this study include Florida Uni-
Costa Rica revealed three distinctive versity,AT&allMahassee (FAMU) and
new species of leptohyphid mayflies, as Texas University, College
described below. Station (TAMU).
VOLUME NUMBER
109, 2 417
Descriptions setae; labial palp three segmented with
Leptohyphes niandibuhis Baumgardner, numerous filiform setae; glossae and
new species paraglossae subequal, fused except distal-
ly, with serrate outer margins; glossae
— (Figs. 1 11) slightly recessed, rounded, and with ro-
Adult. Unkno—wn bust setae; paraglossae with numerous
Mature larva. Body length 3.0- filiform setae. Thorax: Dorsally dark
3.5 mm; caudal filaments 2.0-2.5 mm. reddish brown to black; ventrally pale
General color black. Head: Dark reddish with diffuse black maculae; hindwing
brown to black; small genal projections pads present in males, absent in females.
present; tubercles absent; compound eyes Foreleg: femur (Fig. 4) paleyellow to pale
small and widely separated; three ocelli brown with dorsal, median black spot;
present; antenna light brown, approxi- anterior and posterior margins with scat-
mately two times length ofhead capsule. tered filiform and acuminate setae; dorsal
Mouthparts: Labrum: dorsally with fili- surface with transverse row of elongate
form setae along lateral margin; two setae (Fig. 5). Tibia and tarsus pale
rows of acuminate setae recessed from brown; anterior margin of tibia with 14-
anterior margin; ventrally with one 16evenly spaced, elongate setae; posterior
longitudinal row ofacuminate setae near margin with few, scattered fihform setae;
midline, with interspersed filiform setae; margins of tarsus without setae; ventral
anterior and lateral margins with dense surface with row of four to six robust
filiform and acuminate setae. Right setae; tarsal claw with one submarginal
mandible (Fig. 1): outer and inner in- denticle, and a single row of four to six
cisors two lobed; prostheca arising at marginal denticles, similar in shape and
base of inner incisor with highly size with equal spacing. Mid- and hind
branched, elongate setae projecting to- legs: femorapaleyellowwithdorsal apical
wards molar region; molar region mostly and basal black spots; anterior margins
fused. Left mandible (Fig. 2): outer in- with elongate setae; posterior margins
cisor two lobed, teeth fused almost their with few robust setae (usually on hind
entire distance; inner incisor two lobed; femur) or none; dorsal surface with trans-
prostheca arising at base ofinner incisor, verse rowofrobust setae atbase offemur.
with highly branched setae projecting Tibiae: pale brown, with row of approx-
towards molar region. Hypopharynx: imately 20 evenly spaced elongate setae
lingua apically rounded; numerous fili- along anterior margin; posterior margin
form and acuminate setae present on with few, scattered elongate setae; dorsal
anterior margin; superlinguae oval, with surface with elevated, medio-longitudinal
numerous filiform and acuminate setae ridge. Tarsi pale brown, margins without
along anterior and lateral margins. Max- setae; four to six acuminate setae along
illa (Fig. 3): palp elongate, three seg- inner margin; claws with one submarginal
mented, without apical seta; three sub- denticle, and a single row of four to six
apicalsetaeoninnerapicalmargin;cluster marginal denticles, similar in shape and
of filiform setae on outer apical surface; sizewith equal spacing (Fig. 6). Abdomen:
galealacinia not fused. Labium: submen- Dark reddish brown to black; posterior
tum moderately developed (approximate- margins of terga 1-10 serrate; terga 5-9
ly twice as wide as long), with regularly each with a pair ofelongate setae located
spaced acuminate setae along lateral medially on posterior margin; tubercles
margins; ventrally with numerous robust absent; dorsal lamella of operculate gill
setae most abundant near midline; pre- elongate oval, basal portion black, apical
mentum ventrally with numerous filiform portion pale; scattered acuminate setae
418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-11. Leptohyphesmandibulus, larva. 1, Rightmandible. 2, Left mandible. 3, Maxilla. 4, Foreleg
femur (dorsal view). 5, Foreleg femoral seta. 6, Hind leg claw. 7, Operculate gill (dorsal view). 8,
Operculategill(ventralview). 9,gill 5lamellae(ventralview). 10, Rightmandible(pre-emergentlarva). 11,
Left mandible (pre-emergent larva).
VOLUME NUMBER
109. 2 419
present along inner and apical margins wear associated with feeding. However,
(Fig. 7); ventral lamellae ofoperculate gill numerous larval paratypes associated
transculent-whitish(Fig. 8); gill 5 lamellae with L. mandibulus also displayed this
as in Fig. 9; gill formula: 3/6/6/6/2. Cercus condition, indicating it is probably a nat-
with whorls of robust and elongate setae urally occurring condition resulting from
at each annulat—ion. maturation, and not necessarly a result
Etymology. The specific epithet of of feeding.
this species is a noun from the Latin There does remain the possability that
word mandibula (f), meaning jaw or Leptohyphes mandibulus could be the
mandible. It alludes to the unusual undescribed larval stage of at least one
features ofthe mandibles for th—is species. of five species of Leptohyphes reported
Discussion and diagnosis. The re- from Mexico and Central America,
duced number of outer incisor denticles which were described based upon only
on both the left and right mandibles will the adult stage. These species include
distinguish this species from others in the Leptohyphes berneri Traver (1958) de-
genus Leptohyphes Eaton. While the vast scribed from central Mexico; Lepto-
majority of species within Leptohyphes hyphes brevissimus Eaton (1892) de-
have four denticles on the outer incisor scribed from southern Guatemala based
of the left mandible and three on the upon female subimagos; Leptohyphes
right mandible, L. mandibuhis has only nigripunctum Traver 1943 described from
two outer incisor denticles on each a male subimago from Venezuela and
mandible. In addition, the coloration of later reported from southern Mexico
the operculate gills (basal half dark, (McCafferty 1985); Leptohyphes peter-
apical half pale), and the contrasting seni Ulmer (1920) described from South
coloration of the body and legs also is America based upon male and female
distinct for this species. Leptohyphes subimagoes, and later reported from
cornutus Allen, known only from South Central America (McCafferty 1985); or
America, also has reduced outer incisors Leptohyphes priapus Traver (1958),
similar to L. mandibulus. It can be which was described from Costa Rica.
distinguished from L. mandibulus by the Based upon geography alone, L. priapus
presence of distinct thoracic tubercles, would be the most likely species to be the
which are absent on L. mandibulus. adult stage of Leptohyphes mandibulus.
Also of interest for L. mandibulus are Additional research will be required to
the changes associated with the mand- resolve this situation. —
ibles on mature and pre-emergent larvae. Distribution and biology. This spe-
Mandibular description in the above cies is currently known only from the
species description is based upon rela- type locality in northwestern Costa Rica.
tively mature larvae, but not pre-emer- Other species of leptohyphid mayflies
gent larvae. In pre-emergent larvae, the collected with it included Leptohyphes
outer incisors of the right mandible are zalope Traver and Tricorythodes sordidus
reduced to a single, rounded structure, Allen. —
while the inner incisor is reduced to Type material. Holotype: Mature
a single denticle (Fig. 10). For the left male larva. COSTA RICA: Alajuela
mandible, the outer incisors are fused Province; NE of Bijagua, nr. Las Flores,
into a single, large incisor, while the Rio Areuo (WN1D0°21'06", W85°21'05"),
inner incisor is reduced to a single 07.vi.2000, Shepard (WDS-A-
TAMU.
denticle (Fig. 11). Reduction and fusion 1300). Deposited in Paratypes:
of incisors is very rare among leptohy- Same data as holotype, 6 larvae
phid mayflies and is usually a result of (FAMU), 25 larvae (TAMU).
420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Tricorythodes kirki Baumgardner, regularly spaced acuminate and filiform
new species setae along lateral margins; prementum
(Figs. 12-22) ventrally with numerous filiform setae;
Adult.—Unknown labial palp three segmented with numer-
— ous filiform setae; glossae and paraglos-
Mature larva. Body length 2.5-
3.0 mm; caudal filaments 1.5 mm. Gen- sae subequal, fused except distally, with
slightly serrate outer margins; glossae
eral color reddish brown with extensive
slightly recessed, rounded, and with
black maculae. Head: Pale reddish
robust setae; paraglossae with numerous
brown with variable black maculae on
filiform setae. Thorax: Pale reddish
vertex and posterior to compound eyes;
brown, with extensive black maculae;
tubercles absent; compound eyes small
pro- and mesonota with extensive black
and widely separated; three ocelli pres- shading medially and along margins;
ent; antenna pale, approximately two forewing pads of mature larvae pale
times length of head capsule. Mouth-
brown; hind wing pads absent; filiform
parts: Labrum (Fig. 12): dorsal surface setae present along lateral margin of
with scattered filiform setae; lateral thorax. Femora pale with limited black
margin with numerous filiform and maculation along basal, lateral margin;
acuminate setae; highly branched robust tibiae pale brown with white band at
setae along anterior margin, projecting tibia-tarsus articulation; tarsi pale brown
from ventral surface; ventrally with with median white band; claws pale
longitudinal row ofacuminate setae near brown. Foreleg (Fig. 16): dorsal surface
midline, with interspersed filiform setae; of femur with median transverse row of
lateral margin with numerous filiform filiform and acuminate setae; anterior
and acuminate setae; anterior margin and posterior margins of femur with
with robust, highly branched setae. diffuse filiform and acuminate setae.
Right mandible (Fig. 13): outer incisor Tibia and tarsus: margins with scattered
three lobed, with elongate setae at base acuminate and filiform setae; tarsal claw
of teeth; inner incisor two lobed; pros- (Fig. 17) with single row of 10-12
theca arising at base ofinner incisor with denticles, similar in shape and size with
highly branched, elongate setae project- equal spacing. Mid- and hind legs
ing towards molar region; molar region (Fig. 18): dorsal surfaces of femora with
mostly fused. Left mandible (Fig. 14): few, scattered filiform and acuminate
outer incisor four lobed, mostly fused; setae; anterior and posterior margins
inner incisor two lobed, mostly fused; with numerous filiform and acuminate
prostheca arising at base ofinner incisor, setae. Tibiae: few, scattered filiform and
with branched, elongate setae projecting acuminate setae present along anterior
towards molar region. Hypopharynx: and posterior margins, and dorsal sur-
lingua apically truncate; numerous fili- face. Tarsi: filiform and acuminate setae
form and acuminate setae present on confined to apical margins. Claws
anterior margin; superlinguae oval, with (Fig. 19): with single row of 12 14
numerous filiform and acuminate setae denticles, similar in shape and size with
along anterior and lateral margins. Max- equal spacing. Abdomen: Without tuber-
illa (Fig. 15): palp very short, one cles or elevated carina; terga 1-3 white
segmented, with elongate terminal seta; with black lateral margins; terga 4-6
one or two subapical setae on inner white with limited black meial macu-
apical margin; cluster of dense filiform lation and lateral black margins; terga
setae on outer apical surface. Labium: 7-9 black with numerous filiform setae
submentum moderately developed, with along lateral and posterior margins;
VOLUME 109, NUMBER 2 421
Figs 12-22. Tricorvhvphes kirki. larva. 12, Labrum (left: dorsal; right: ventral). 13, Right mandible.
14, Left mandible, is'. Maxilla. 16, Foreleg. 17, Foreleg claw. 18, Hindleg. 19, Hind leg claw. 20,
Operculate gill (dorsal view). 21, Operculate gill (ventral view). 22, Gill 3.
—
422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tergum 10 brown. Dorsal lamella of north-central Costa Rica. Larvae were
operculate gill (Fig. 20) on abdominal collected from a very small, shallow first
segment 2 triangular with margins or second order stream at elevations
rounded; narrow, elongate ridge present ranging from 200 to 650 m. Substrate
near inner margin; basal one-halfto two- of the steams was composed mostly of
thirds of gills covered with black macu- rock-rubble with little coarse gravel.
lae; apical margin pale; acuminate and Other mayflies collected at these sites
filiform setae present along entire outer included Tricorythodes sordidus Allen
margin; ventral lamellae of operculate (Leptohyphidae) and Camelobaetidius
gill pale translucent (Fig. 21); gill 3 as in warreni (Traver and Edmunds) (Baeti-
figure 22; gill formula: 2/3/3/3/2. Cercus dae).
with whorls ofelongate and robust setae Type material.—Holotype: Mature
at each annula—tion. male larva. COSTA RICA: Alajuela Pro-
Etymology. This species is named for vince, Rio Guayabo at Hwy 140, 1.8 km
my son, Kirk, in recognition of his E Venicia (N10°40'45N, W84"15'13", elev.
support and assistance. — 460 m), 09.vi.2001, DE Baumgardner
Discussion and diagnosis. Only three (DB 01-25). Deposited at TAMU. Para-
other species of Tricorythodes Ulmer are
currently known from Central America, types: Same data as holotype, 5 lar-
south of Mexico. The common and vae 0$, 2$), 2 slides (DB05012801,
DB05xi2208) (4 larvae and slides TAMU,
pwriidmeulsy,-dni.stsrpi.b(usteeeddiTs.crsioprtdiiodnubselAolwl)e,n,anTd. 1 larva FAMU). COSTA RICA: Heredia
T. costariccmus (Ulmer), known only PrKovmince, unnamed creek at Hwy. 4, ca.
from the adult stage. Tricorythodes kirki 3 fromjet. with Hwy. 32 (N10°15'10",
larvae can be differentiated from T. W83°55'll"; elev. 200 m) 10.vi.2001, DEB
sordidus larvae by the lack of elongate (DB 01-28), 22 larvae (TAMU); La Selva
SW
filiform setae on the body, which are Biological Station, Puerto Viejo, Sura
present on T. sordidus. In addition, T. Creek at Rio Puerto Viejo (N10°25'49";
sordidus larvae possess abdominal terga W84°00'06", elev. 33 m), 09.vi.2001, 8L,
which are uniformly pale brown in color, DEB (DB 01-26), 8L (5L TAMU, 3L
while T. kirki larvae have contrasting FAMU); Rio Isla Grande at Hwy. 4, ca.
coloration of the abdominal terga as W5 Km. W. of Rio Frio (N10°23'31";
described above. Tricorythodes kirki lar- 83°58'04", elev. 65 m), 10.vi.2001,
vae can be distinguished from larvae of DEB (DB 01-27), IL (PERC). San Jose
T. pirmus by the presence ofa distinctive Province; Rio Pedregoso at Hwy. 243, ca.
Km
and well-developed frontal shelf and 4 S. San Isidro de El General (N09°
genal projections, which are absent in 21T5 W83°43'35", elev. 660 m), 22.vi.
T. kirki. There does remain the poss- 2001, DEB (DB 01-47), 12L [TAMU].
ability that T. kirki could be the unde-
scribed larval stage of T. costaricanus. Tricorythodesprimus Baumgardner,
However, considering the very dark new species
brown coloration of T. costaricanus
(Figs. 23-31)
adults, and the very limited dark color- —
ation of mature T. kirki larvae, the Adult. Unknow—n.
possability appears remote that the two Mature larva. Body length 3.0-
are indeed the same species. 3.5 mm; caudal filaments 3.0 mm.
Distribution and biology. Tricor- General color yellow with black maculae
ythodes kirki is known only from Ala- on thorax and abdomen. Head: (Fig. 23)
juela, Heredia, and San Jose provinces in yellow without black maculae; large
VOLUME NUMBER
109, 2 423
Figs. 23-31. Thcoyrhyphesprimus, larva. Fig. 23, Head and pro- and mesothorax (dorsal view). Fig.
24, Right mandible. Fig. 25, Leftmandible. Fig. 26, Maxilla. Fig. 27, Labium(left: dorsal; right: ventral).
Fig. 28, Foreleg. Fig. 29, Foreleg claw. Fig. 30, Operculate gill. Fig. 31, Operculate gill (ventral view).
424 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
genal and frontoclypeal projections pres- pair of distinctive, sharp projections on
ent; tubercles absent; compound eyes anterior lateral margin (Fig. 23); lateral
small and widely separated; three ocelli margins with scattered acuminate setae;
present; margins of head with filiform mesonotum (Fig. 23) yellow with exten-
setae; antenna pale, approximately as sive and irregular black maculation, and
long as head capsule. Moiithparts: La- scattered acuminate setae along lateral
brum: dorsally with filiform setae along margins. Legs: all segments of all legs
lateral margin; acuminate setae recessed pale yellow. Foreleg (Fig. 28): dorsal
from anterior margin; ventrally with one surface of femur with a transverse row
longitudinal row ofacuminate setae near of numerous filiform setae; anterior and
midline, with interspersed filiform setae; posterior margins with numerous acumi-
anterior margin with filiform setae. nate and filiform setae. Tibia and tarsus:
Right mandible (Fig. 24): outer incisor anterior and posterior margins with
three lobed; inner incisor two lobed; numerous acuminate and filiform setae.
prostheca and molar region as in Claw (Fig. 29): with two or three, minute
Fig. 17; numerous and very long, thick marginal denticles, and two pair of
setae present along outer margin of submarginal denticles. Mid- and hind
mandible. Left mandible (Fig. 25): outer legs: dorsal surfaces of femora with
incisor four lobed, mostly fused; inner numerous, irregularly spaced filiform
incisor two lobed; prostheca arising at and acuminate setae; anterior and pos-
base of inner incisor, with filiform setae terior margins with numerous acuminate
projecting towards molar region; numer- and filiform setae. Tibiae: filiform setae
ous and very long, thick setae present present along anterior and posterior
along outer margin of mandible. Hypo- margins. Tarsi: margins with numerous
pharynx: lingua apically truncate; nu- filiform setae. Claws: with two minute
merous filiform and acuminate setae denticles and two pair of submarginal
present on anterior margin; superlinguae denticles. Abdomen: Pale yellow with
oval, with numerous filiform and acumi- dorsal and ventral transverse black
nate setae along anterior and lateral bands on segments 1-9; posterior mar-
margins. Maxilla (Fig. 26): palp one gins of terga 1-9 with numerous filiform
segmented, with terminal seta; two sub- setae; lateral margins of segments 2-7
apical setae on inner apical margin; expanded outward with filiform setae
cluster of filiform setae on outer apical present along margin. Dorsal lamella of
surface. Labium (Fig. 27): submentum operculate gill (Fig. 30) on abdominal
moderately developed, with regularly segment two triangular, pale yellow with
spaced acuminate and filiform setae basal black band at base, and very sparse
along lateral margins; ventrally with black maculae over dorsal surface; fili-
numerous robust setae; prementum ven- form setae present along entire margin;
trally with numerous acuminate setae; ventral lamellae of operculate gill pale
labial palp three segmented with numer- translucent (Fig. 31); gill formula: 1/3/3/
ous filiform setae; glossae and paraglos- 3/2. Cercus with whorls of elongate and
sae subequal, fused except distally, with robust setae a—t each annulation.
slightly serrated margin; glossae serrated Etymology. The specific epithet ofthis
with robust setae along margin; para- species is a noun from the Latin word
glossae with numerous filiform and primus (m), meaning first or original. It
acuminate setae. Thorax: (Fig. 23) yel- refers to the first description ofa species of
low with extensive black maculae; pro- Tricorythodes from Central America which
notum with a pair ofblack inverted "L" possesses a distinctive and well-developed
shaped markings lateral to midline, and frontal shelf and genal projections.
——
VOLUME NUMBER
109, 2 425
Discussion and diagnosis. Tricor- ince; Rio Caracol at CA Hwy. 2, ca.
Km
ythodes primus is most similar to T. 7.3 E. Rio Claro (N08°39'47",
condylus Allen from southwestern Unit- W83°00'41", elev. 80 feet), 23.vi.2001,
ed States, and T. popyanicus Dominguez DE Baumgardner (DB 01-55). Deposited
from Central America. Tricorythodes in TAMU. Paratypes: COSTA RICA:
primus can be distinguished from T. Puntarenas: Rio Coloradito at CA
condylus by its much smaller size (3.0- Hwy. 2, ca. 6.7 Km SE Ciudad Neily
mm mm
3.5 for T. primus versus 5.0-6.0 (N08°36'09", W82°52'02", elev. 180 feet),
for T. condylus) and more extensive 23.vi.2001, DE Baumgardner (DB 01-
black maculation of the thorax and 56), 1 larva (legs and mouthparts
abdomen. It can be distinguished from mounted on slide #05x12006), slide and
T. popyanicus by the presence of well- specimen deposited in TAMU.
developed antero-lateral projections of
Acknowledgments
the pronotum, which are very weakly
developed on T. popyanicus. In addition, I thank the Institutio Nacional de
T. popyanicus is known only from Biodiversidad, San Jose, Costa Rica,
southern South America while T. primus for arranging collecting permits and
is known only from Central America. assisting with travel while in Costa Rica,
Tricorythodes popyanicus and T. con- and W. D. Shepard for donation of
dylus (along with three other species) specimens. Funding for field work in
were recently transferred to the genus Costa Rica was provided by the L.T.
A&M
Tricoryhyphes (Wiersema and McCaff- Jordan Fellows Program ofTexas
erty 2000), which was originally de- University. Thanks also to Boris Kon-
scribed by Allen and Murvosh (1987) as dratieff (Colorado State University),
A&M
a subgenus of Tricorythodes, and elevat- Wills Flowers (Florida Universi-
ed to generic status by Wiersema and ty), Carlos Molineri (National Universi-
McCafferty (2000). Molineri (2002) ty of Tucuman, Argentina) and W.P.
clearly showed Tricoryhyphes to be McCafferty (Purdue University) for re-
synonymous with Tricorythodes, synon- viewing an earlier draft of the manu-
ymizing the latter with the former. Un- script.
published data by D.E. Baumgardner
also supports the conclusions ofMolineri Literature Cited
(2002). Allen, R. K. and C. M. Murvosh. 1987. Mayflies
Distribution and biology. Tricor- (Ephemeroptera: Tricorythidae) of the South-
ythodes primus was collected from clear western United States and Northern Mexico.
Annals ofthe Entomological Society ofAmer-
flowing streams which were several me-
ica 80: 35-^0.
ters wide, shallow, and with substrate Baumgardner, D. E. 2003. New synonyms and
varying from sand to boulders. These low stage description for three species ofLeptohy-
elevation costal streams are located in the phidae (Ephemeroptera). Proceedings of the
extreme southeastern region ofthe coun- Entomological Society of Washington 105:
try, near the Panama border . Other Baum2g0a3r-d2n0e8.r, D. E. and Socorro Avila, A. 2006.
species of mayflies collected with T. Cahecar serratus, a new genus and species of
primus included Thraulodes brunneus leptohyphid mayfly from Central America,
Koss, Thraulodes spp. (Leptophlebiidae), and description of the imaginal stages of
LeptohypheszcdopeTraver, Tricorythodes Tricorythodes sordidus Allen (Ephemeroptera:
sordidus Allen, and Vacupernius packeri Leptohyphidae). Zootaxa 1187: 47-59.
Baumgardner, D. E. and W. P. McCafferty. 2000.
(Allen) (Leptohyphi—dae). Leptohyphes zalope (Ephemeroptera: Leptohy-
Type material. Holotype: Mature phidae): a polytypic North and Central Amer-
larva. COSTA RICA: Puntarenas Prov- ican species. Entomological News 111: 49-59.