Bijdragen tot de Dierkunde, 63 (3) 173-191 (1993)
SPB Academie Publishing bv, The Hague
New records and redescriptions of American species of Mesocyclops and
of Diacyclops bernardi (Petkovski, 1986) (Copepoda: Cyclopoida)
Janet W. Reid
Department of Invertebrate Zoology, NHB-163, National Museum of Natural History, Smithsonian
Institution, Washington, DC 20560, U.S.A.
Keywords: Taxonomy, Copepoda, Cyclopoida, Mesocyclops, new records, U.S.A.
Abstract Introduction
Recent collections in the south central U.S.A. have included Taxonomie understanding of the tropico-temperate three neotropical and one probably introduced species that are has eyelopoid eopepod genus Mesocyclops greatly presently assigned tothe cyclopoid copepod genus Mesocyclops. improved during the past decade. Recognition of
Mesocyclops longisetus var. curvatus Dussart, 1987, is reported the importance of formerly ignored subtle morpho- from Louisiana, U.S.A., and Panama; published records of logical usually meristic characters by Kiefer( 1981), Mesocyclops longisetus (Thiébaud, 1914) sensu lato in the ,
Van de Velde and Dussart & Fernando southern U.S.A. are reviewed. Mesocyclops reidae Petkovski, (1984a, b),
is reported from Mississippi, U.S.A., and Honduras. 1986, (1988) has led to more accurate characterization of Mesocyclops ruttneri Kiefer, 1981, is redescribed from type many paleotropical species. It is desirable to supply specimens collected in Austria, and newly reported from similarly detailed descriptions for the American Louisiana, Mississippi, China, Thailand, and Viet Nam. This species. species is considered to have been introduced from Asia into
Austria and the U.S.A. Mesocyclops bernardi Petkovski, 1986, New records of copepods from non-lacustrine newlyrecorded from Louisiana and Mexico, is transferred to the wetlands in the southern U.S.A. have extended the
genus Diacyclops. known geographical ranges of many species. Neo-
tropical species appear to comprise a significant
and previously little appreciated component of the Résumé regional copepod fauna (Reid, 1992). Several neo-
tropical species appeared in collections of cyclopoid
Dans des prélèvements récemment réalisés au sud des parties copepods that were made from Louisiana and Mis- centrales des U.S.A. ont été découvertes trois espèces néo- sissippi, U.S.A. by Gerald G. Marten and sent to tropicales et une espèce probablementintroduite de Copépodes me for determination 1989; 1990a, In Cyclopoïdes actuellement considérés comme appartenant au (Marten, b).
Dus- addition to the North American genre Mesocyclops. Mesocyclops longisetus var. curvatus common Mesocy- sart, 1987, est mentionné de Louisiane (U.S.A.) et de Panamá; clops edax(S.A. Forbes, 1891), there were four spe- sont passés en revue les mentions publiées de Mesocyclops cies that are the same presently assigned to genus: longisetus (Thiébaud, 1914) sensu lato dans les zones méri- Mesocyclops bernardi Petkovski, 1986, Mesocy- dionales des U.S.A. Mesocyclops reidae Petkovski, 1986, est
var. curvatus Meso- connu du Mississippi (U.S.A.) et du Honduras. Mesocyclops clops longisetus Dussart, 1987,
reidae and ruttneri Kiefer, 1981, est redécrit sur des exemplaires-type cyclops Petkovski, 1986, Mesocyclops
mentionné Missis- d’Autriche, et nouvellement de Louisiane, du ruttneri Kiefer, 1981. Examination of material in sippi, de Chine, Thaïlande, et Viet Nam; on considère cette es- the C.D. Marsh Collection of Copepoda in the pèce comme ayant été introduite d’Asie en Autriche et aux National Museum of Natural History led to new U.S.A. Mesocyclops bernardi Petkovski, 1986 — nouvellement
recordsof M. longisetus var. curvatus andM. reidae mentionné pour la Louisiane et le Mexique — est transféré au
from Panama and genre Diacyclops. Honduras, respectively. The 174 /. W. Reid - New records of American Mesocyclops and Diacyclops bernardi
present article describes aspects of the morphology Discussion. — The New Orleans specimens are con-
the the and reviews the known distribution of M. longise- gruent in proportions of caudal rami, the
form of the antennular and in the tus var. curvatus, M. reidae, and M. ruttneri. The membrane, leg 4
3, with M. var. transfer of M. bernardi to the genus Diacyclops is endopodite longisetus curvatus
1987. The lateral of the seminal proposed. An additional new record for D. ber- Dussart, arms
of nardi is given from Mexico. receptacles most specimens examined are strong-
recurved In a few these For taxonomie examination, specimens were ly posteriorly. specimens,
little recurved therefore closer treated according to the methods of Reid et al. arms are posteriorly,
M. restricto (1989). Terminology, particularly for the seminal to longisetus (Thiébaud, 1914) sensu receptacle and the antennule is taken from Fiers & Dussart, 1987.
The record from New Orleans was previously Van de Velde (1984) and Van de Velde (1984a, b). reported by Marten (1989, 1990b). The earliest Descriptions of the morphology of species of
record of M. longisetus sensu lato in the U.S.A. Mesocyclops are based on the redescription of the Herrick may be that of (1887), whose extensive genotype, M. leuckarti (Claus, 1857) by Van de figures of “Cyclops simplex” from southern Velde(1984a). Specimens are deposited eitherin the
Alabama clearly show a species of Mesocyclops collections of G.G. Marten, New Orleans Mosquito
with the caudal ramus haired on the medial surface, Control Board, or in the U.S. National Museum of
theantennularhyaline membrane with a single deep Natural History, Smithsonian Institution (USNM). notch, and the leg 4 coupler with two triangular
projections. However, the illustration of the semi-
nal receptacle (Herrick, 1887: plate VII, fig. la) Taxonomic section shows the medianthird of the anterior margin con-
cave and the lateral arms tapering, not rounded as Family Cyclopidae Burmeister, 1834 the receptacle of M. longisetus. Reddell (1965, Subfamily Cyclopinae Dana, 1853, char, emend. repeated in Reddell & Mitchell, 1969) reported M. Kiefer, 1927 longisetus from Felton Cave, Sutton County, Genus Mesocyclops G.O. Sars, 1914 Texas. The species was also recorded from Florida
by Dussart & Fernando (1986), but additional local-
data for their record ity are unavailable (C.H.
var. curvatus 1987 Mesocyclops longisetus Dussart, Fernando, in litt., 1990). It seems that M. longise-
tus sensu lato, which is distributed through Mexico
- and Central and South America as far south as Synonymy. Mesocyclops longisetus var. curvatus Dussart,
1987: 150, 156, figs. 3, 4, 7, 8; Reid, 1990: 181, table I. Patagonia and possibly Tierra del Fuego, occurs
Mesocyclops longisetus (Thiébaud, 1914); Marten, 1989: 232, widely although sporadically in the southernU.S.A.
233, 235, table 1; 1990b: 681-687, tables 1,4, 5, 7, 8; Reddell, The distribution of M. longisetus var. curvatus is 1965: 157; ?Reddell & Mitchell, 1969: 7; ?Dussart & Fernando, uncertain. 1986: 291, 292. The records for Panama from the Marsh Collec- Cyclops simplex Poggenpol, 1874; ?Herrick, 1887: 14, 17-18, tion pl. VII fig. 1. are previously unpublished.
Material. - USNM 252008, 699, samples Ml and M6, Lake Mesocyclops reidae Petkovski, 1986 Catherine marshes, 24 July 1991, after introduction from cul- tures originally collected from Joe Brown Lagoon, New (Figs. 1, 2)
coll. G.G. Orleans, Louisiana, Marten. One 9, dissected on slide, and 2 9 9, ethanol-preserved, New sample 105, Orleans, Synonymy. - Mesocyclops reidae Petkovski, 1986: 47, 61-66,
Louisiana, 1988,G.G. Marten collection. USNM Accession No. 71-73, 78, Abb. 11-22; 1988: 40; Sket, 1988: 79, table 1; Reid, 120079 (Marsh Collection), several 9 9 on slides, Panama, 1990: 181, table 1.
C.D. Marsh prep. nos. 3826, 3827, 3842, 3843, and 3943. Un- Mesocyclops ellipticus (non Kiefer, 1936a); Yeatman, 1977: mounted in 70% specimens preserved ethanol. 5-7, figs. 1-16; Smith & Fernando, 1978: 2015, 2016, 2019, Bijdragen tot deDierkunde, 63 (3) - 1993 175
Fig. 1. Mesocyclopsreidae Petkovski, 1986, dissected from Mississippi, USNM 252009: a, pediger 5 and genital segment, right lateral;
setae not and b, pediger 5 and genitalsegment, ventral; c, left leg 5, lateral-oblique, completelyindicated; d, copulatorypore pore-canal, lateral-oblique;e, anal somite and caudal rami, dorsal; f, anal somite and caudal rami, ventral; g, antennule articles 16 and 17; h, anten-
indicated Scale to na article 1, caudal side; i, antenna article 1, frontal side; j, labrum, spines only on one side; k, maxilliped. applies
Figs. 1a-c only. 176 J. W. Reid New records of American Mesocyclops and Diacyclops bernardi
2020, figs. 21-24, tables 1, 2; 1980: 11, 18,20, fig.9A-D, table margin (indicated by arrow) short, no other surface
2; Reddell, 1981: 81; Pesce, 1985: 296, 311, 316-318, figs. ornamentation.
63-65. of Labrum (Fig. lj) with 2 rows long spines on
each side of median line. Maxillule as in M. leu-
Material. - USNM 252009,1 9 , dissected onslide, and 10 9 9, ckarti, i.e. lacking spines on surface of palp. Maxil- ricefield, Cleveland, Mississippi, September 1991, coll. G.G. la in M. surface of Marten. USNM Accession No. 120079 (Marsh Collection), 2 as leuckarti, lacking spines on
9 9 on slides, C.D. Marsh prep. nos. 4288, 4296, Honduras. coxa (article 2), distal half of ventral surface of
coxa rugose. Maxilliped (Fig. Ik) article 2 (basis)
Description of female. - Range of lengths of Mis- with 3 transverse rows of small spines along poster-
sissippi specimens 0.80-1.02 mm (median = 0.90 ior border; middle seta of article 4 (endopodite 2)
mm, n = 10). Resembling description of Petkovski short in comparison to corresponding seta of con-
(1986) of type population from San Andrés Island, geners.
Colombia, in most details. Legs 1-4 as in M. leuckarti except in following
Supplementary observations: Posterolateralmar- details. Leg 1 (Fig. 2a) with medial seta of basipo-
gin of pediger 4 finely serrate (Fig. la). Pediger 5 dite short, stout, with spines at base; coupler with
(Figs, la, b) with small lateral spines disposed in 2 rounded marginal expansions. Leg 2 (Fig. 2b),
3 diagonal rows. Leg 5 (Figs, la-c) with median medial expansion of basipodite haired, coupler
inserted distal } of article 2. Genital with 2 blunt spine at A seg- large triangular expansions. Leg 3 (Fig.
ment medial of (Figs, la, b) lacking ornamentation except 2c), expansion basipodite haired, expan-
sions few papillae bearing sensilla near posterior margin. of coupler large, broadly triangular, acute.
Pore-canal of seminal receptacle (Figs, lb, d) Leg 4 (Figs. 2d, e), posterodistal margin of coxopo-
short, directed dorsally, in semilateral view seen dite with short row of small spines; basipodite
to be recurved anteriorly. Copulatory pore with naked medially; coupler with 2 medially directed
slightly sclerotized anterior margin. Anal somite slenderacuminate spiniform processes; and most of
(Figs, le, f), posterodorsal margin lacking spines; lateral margin of medial terminal spine of endopo-
postero ventral margin with small spines, mediad dite article 3 finely spinulate.
spines larger. Caudal ramus (Figs, le, f) similar to
of Petkovski Abb. in - description (1986: 62, 13) Discussion. Principal characters such as the
lacking spines near base of lateral seta, but having characteristic marginal protrusions of the swim-
2 tiny spines anterior to base of lateralmost termi- ming leg couplers, the short stout medial spine of
nal caudal seta in both Mississippi and Honduras the leg 1 basipodite, and the relative lengths of the
specimens. Medialmost terminal caudal seta less caudal setae with the agree so closely ample descrip-
than twice of lateralmost terminal caudal length tion of Petkovski (1986) that I have no hesitationin
seta. assigning the specimens from Mississippi and Hon-
Antennule article small in 1, spines disposed row duras to this taxon. Petkovski (1986: 64, Abb. 18)
in M. articles as leuckarti; 2-17 lacking surface did not observe spinules on the lateral margin of the
of antennule in spines; appendages exactly as leg 4 endopodite 3 medioterminalspine, but these
M. leuckarti (Van de Velde, 1984a: 14, fig. 5A). are very fine and difficult to see in some of the
Antennule membranes hyaline (Fig. lg), membrane specimens at hand.
of article 16 rather narrow, margin finely serrate Both the Honduras and Mississippi records are
than entire as observed Petkovski by (1986: 62, previously unpublished. Petkovski (1986) syn-
Abb. of 14); membrane article 17 broader, slightly onymized several records of M. ellipticus with his
more from coarsely serrate, extending distally seta new species M. reidae. With the discovery of M. rei-
near midlength of article. dae in Mississippi the known range of this species
Antenna with (Figs, lh, i) basic spine pattern is extended significantly northward from Central
for genus, spine rows, especially row on anterior America, Mexico, and the Antilles. Bijdragen tot de Dierkunde, 63 (3) - 1993 177
2. 1 Fig. Mesocyclops reidae Petkovski, 1986, dissected from Mississippi, USNM 252009: a, left leg and coupler, anterior, most of rami not indicated; b, left leg 2 coxa-basipodite and coupler, anterior; c, left leg 3 coxa-basipoditeand coupler, anterior; d, right leg
4 4 and coupler, posterior, most of rami not indicated; e, right leg endopodite article 3, posterior, setules of setae not indicated.
Mesocyclops ruttneri Kiefer, 1981 Material. - Paratypes from the F. Kiefer Collection,Staatliches
(Figs. 3-5) Museum fiir Naturkunde Karlsruhe: Mikropráparat 11101, 2
9 9 Abd. + P5, prep. Kiefer, 30 September 1980; Mikroprá-
1 parat 11102, 9 A1-P4, prep. Kiefer, 30 September 1980;
11 - Mikropráparat several 9 9 Synonymy. Mesocyclops ruttneri Kiefer, 1981: 151,156, 178 — 11285, Al, 4A2, prep. Kiefer,
November than 20 Glas No. from 180, 186, 187, Abb. 1 (54, 55), 14; Marten, 1990b: 681-688, 1982; more adults, 470; 1 9
Glas No. dissected all from Warm- tables 1-8. 470, on slide, prep. Reid;
wasser im Glashaus, Lunz, Österreich, 4 December 1926, coll. Mesocyclops sp. (leuckarti-group); Marten, 1989: 232-235,
Klie? mark Kiefer's table 1; 1990a: 160. (question according to label).
Non M. pehpeiensis (Hu, 1943); Dussart & Fernando, 1985:
246; 1988: 249; Lim & Fernando, 1985: 73, 80, 83-85, figs. Additional, non-paratype material: USNM 90830, 24 ç ç,
57-59. ponds, Foochow, Fukien Province, China, April 1948, coll.
?Mesocyclops Leuckarti pehpeiensis Hu, 1943: 115, 124-126, C.C. Tang. USNM 250683,1 Ç 2 o- 8 Lumtak- , er, copepodids, table II, fig. c. long Creek, Khao Yai National Park, 100 km northeast ofBang- 178 J. W. Reid New records of American Mesocyclops and Diacyclops bernardi
3. Fig. Mesocyclops ruttneri Kiefer, 1981, paratype from the Kiefer Collection (a, d, e, from Glas 470; b, c, from Mikropräparat
from 5 and somite 11101; f, g, Mikropräparat 11102):a, habitus, dorsal; b, pediger genital segment, ventral; c, anal and caudal rami, dorsal; antennule article article caudal article frontal side. Scale d, antennule, setae not indicated; e, 17; f, antenna 1, side; g, antenna 1, applies to Fig. 3a. - Bijdragen tot de Dierkunde, 63 (3) 1993 179
kok, Thailand, 24 January 1989, coll. T. Ishida. USNM 250565, to insertion of lateral seta, and 5 or 6 larger spines
ricefield New Loui- 699, on Compton Farm, near Orleans, dorsal and lateral to base of lateralmost terminal
siana (after experimental introduction), 10 August 1990, coll. caudal seta; medial surface of ramus naked. Most
G.G. Marten. USNM 252010, 2 9 9, each dissected on slide, caudalsetae with fine uniformplumage, dorsal seta Lagoon (canal) in Joe Brown Park, east New Orleans, Loui-
naked. of of caudal setae as in siana, July 1987, coll. G.G. Marten. USNM 252011, 1 9,8 Proportions lengths
crcr, 5 copepodids, sample No. 70, Lagoon (canal) in Joe Fig. 3a.
Brown Park, east New Orleans, Louisiana, 10 June 1988, coll. Antennule (Figs. 3a, d, e) when completely
G.G. Marten. USNM 252012, 1 9, dissected on slide, and 28 reflexed reaching slightly past posterior margin of 9 9, sample No. 105, Lagoon (canal), Joe Brown Park, east pediger 2, setation like that of M. leuckarti; articles New Orleans, Louisiana, 30 June 1988, coll. G.G. Marten. and 7-13 with of of small 4, 5, rows spines, USNM 252013, 23 adult specimens, culture,originallyfrom east groups
New Orleans, Louisiana, 1989, coll. G.G. Marten. USNM with some variation:article 5 with 2 rows of spines
252014,14 , 6 a- 75 copepodids,samples 209-210, from 9 9 a, on most specimens, withadditionalrow of 3 spines field trials, New Orleans, Louisiana, 1989, coll. G.G. Marten. on 1 antennuleof female on Mikropràparat 11102. USNM 250564, 1 9 ricefield, Bebe near Jennings,Loui- , Farm, Antennules of some specimens with few shallow siana, 9 August 1990, coll. G.G. Marten. USNM 264729,4 9 9, round pits on dorsal surface of article 1. Hyaline True Bach Lake, Hanoi, Viet Nam, 12 January 1993, coll. Vu
Sinh Nam. G.G. Marten collection: more than 20 9 9, Jennings membranes of antennule articles 16 and 17 finely
26 and than 20 9 ricefield, July 1991; more 9, ricefield, serrate, membrane of article 17 with deep notch.
Cleveland, Mississippi, 10 August 1991. Undissected specimens Antenna(Figs. 3f, g) general structure including ethanol-preserved. setation like that of M. leuckarti; article 1, in addi-
tion to basic spine pattern of genus (Van de Velde,
Description of female. — Length as given by Kiefer 1984a, b), caudal side with oblique row of tiny
(1981) for type population, 1.0-1.15 mm; lengths spines near medial margin, single or doubleirregu-
of 10 unmounted paratype specimens, 1.0-1.28 lar transverse row of 6-14 larger spines at level of
of from 1.10-1.23 medial dis- mm; specimens Fukien, China, proximalmost seta, and 2-4 spines near
= mm (median - 1.16 mm, n 10); of specimen tal margin.
from Thailand, 1.26 mm; of specimens from Labrum, mandible, maxillule, and maxilla as
Compton Farm, Louisiana (USNM 250565), 0.98— corresponding structures of M. leuckarti. Maxil-
1.19 mm (median = 1.04mm, n = 6); of specimens liped (Fig. 4a) with row of small spines on article 1
from Viet Nam, 1.34-1.56 mm (median = 1.41 and 2 groups of many small spines on article 2;
mm, n = 4). setules of most setae sparse.
of Habitus medial of Redescription paratype specimens: Leg 1 (Fig. 4b), expansion basipodite
(Fig. 3a) slender; pediger 5 little expanded laterally, lacking seta, leg otherwise similar in setation and lacking ornamentation on lateral and dorsal sur- ornamentationto leg 1 of M. leuckarti. Legs 2 and faces similar of leuckarti except two hairs near dorsomedian line as 3 to those M. except lacking
of present in all congeners. Genital segment (Figs. 3a, group tiny spines near proximolateral corner of b), anterior halflittleexpanded, few scattered shal- posterior surface of coxopodite, present in M. low circular pits on lateral and dorsal surface of leuckarti (location indicated by arrow in Fig. 4c).
surface similar that of posterior half, including area posterior to Leg 4 (Figs. 4d-h) to M. leuckarti, leg 6 otherwise without ornamentation. Seminal except medial expansion of basipodite naked and receptacle (Fig. 3b) with anterior margin slightly coxopodite with 2 transverse rows of spines along
well concave, lateral arms nearly horizontal, their later- posterodistal margin, these rows separated in al of ends slightly recurved anteriorly; posterior mar- most specimens; 1 or 2 rows small spines on gins leading anteriorly from copulatory pore, more posterior surface of coxopodite near lateral margin or less fused near pore in differentspecimens before variously developed in different specimens, these
curved in diverging laterally; pore-canal long, posteri- rows lacking Louisiana specimens (Figs. 4d, e, orly. Caudal rami (Figs. 3a, c) about 3.2 times arrows). Marginal spiniform processes of leg 4 longer than broad, bearing 1 or 2 tiny spines dorsal coupler (Figs. 4d, g, h) more or less triangular, 180 /. W. Reid - New records of American Mesocyclops and Diacyclops bernardi
4. ruttneri the Kiefer Collection from Glas Fig. Mesocyclops Kiefer, 1981, paratype from (a, b, d 470; c, e, f, g, h from
h from different 1 Mikropräparat 1102; g, specimens): a, maxilliped; b, leg and coupler, anterior; c, leg 2 coxa-basipodite, posterior;
4 and d, leg coupler, posterior; e, leg 4 coxa-basipodite, posterior; f, leg 4 endopodite article 3; g, h, leg 4 couplers. Bijdragen tot de Dierkunde, 63 (3) - 1993 181
Fig. 5. Mesocyclops ruttneri Kiefer, 1981, a-c, dissected from Louisiana,USNM 252012; d, e, paratype from the Kiefer Collec-
antenna tion, Glas 470: a, antennule, setae omitted; b, antenna article 1, caudal side; c, article 1, frontal side; d, habitus, dorsal; e,
pedigers 5, 6 and succeeding urosomite. Scale applies to Fig. 5d.
always with acute curved slender tip. Leg 5 (Fig. ably constant in the individuals and populations
3b), seta of article 1 shorter than seta of article 2, examined. The variationof antenna spine patterns
distant seta and spine of article 2 subequal in length. (Figs. 3f, g, 5b, c) between populations was
The antennulespine patterns (Figs. 3d, 5a) varied no greater than the within-population variation.
somewhat in the number and arrangementof spines The shape of the spiniform processes of the leg 4
varied from trian- on each article, but the general pattern was remark- coupler (Figs. 4d, g, h) bluntly - 182 J. W. Reid New records of American Mesocyclops and Diacy clops bernardi
guiar to very slender, usually similar to Fig. 4d. not indicated) and Lim & Fernando (1985, Malay-
sia) for the caudal side of antenna article 1 of “M.
Description of male. — Lengths of paratypes from pehpeiensis” show alarge diffuse group rather than the Kiefer Collection Glas No. 470, 0.73-0.89 mm a row of spines at the level of the proximal medial = Habitus (median 0.82 mm, n = 10). (Fig. 5d) distal seta, and no spines near the margin. The two slender. Antennule geniculate, setation and aesthe- of level distal to the groups tiny spines at a longitu- tascs of articles 1-4 similar to that described for dinalrow of spines on the frontal side of this article male of M. leuckarti by Gurney (1933: 290, fig. shown by Dussart & Fernando (1986) and Lim & 1865). Dorsal surface of antennule article 1 of all Fernando in M. ruttneri. (1985) are not present j specimens examined with several transverse rows of Seminal receptacles also differ: in M. ruttnerithe round pits. Mouthparts and legs 1-5 as in female. lateralarms are narrower and slightly recurved an- Leg 6 (Fig. 5e) composed of small trapezoidal flap teriorly at the lateral ends. The posterior margins, bearing 1 stout ventral spine, 1 short median seta, from the in leading copulatory pore, M. ruttneriare and 1 long dorsal seta reaching posterior margin of always directed anteriorly and usually conjoined to succeeding somite. some extent from the pore before diverging lateral-
Discussion. - Specimens of Mesocyclops ruttneri ly in a sharp curve, while in all representations of
collected in Austria in were first a greenhouse, “M. pehpeiensis” these margins curve laterally which had been destroyed by the time thatthe origi- directly from the pore. nal description was published (Kiefer, 1981). Kiefer In M. ruttneri there are always one or two small
(1981) speculated that the species must be Asian. spines on the caudal ramus proximal to the inser-
Dussart & Fernando (1985, 1988) and Lim & tion of the lateral caudal seta. Spines at this loca-
Fernando (1985) described a species from Austra- tion are shown by neither Dussart & Fernando
and Sri Lanka Lim Fernando lia, Burma, Indonesia, Malaysia, (1988) nor & (1985). and identified it as M. pehpeiensis Hu, 1943, a Finally, Lim & Fernando (1985) show the medial poorly known species originally described from expansion of the leg 4 basipodite as thickly haired.
China. They synonymized M. ruttneri with M. peh- Hu's (1943) original description of M. pehpeien- peiensis on the basis of similarities betweenKiefer's sis provides few of the morphological details that
and their without considered to description material, apparently are today necessary distinguish spe- comparing type material of M. ruttneri. These cies of the genus. Hu gave the lengths of female authors' figures of various populations assigned to specimens as 1.525-1.710 mm. He reported the
M. pehpeiensis differ among themselves in several caudal ramus of M. pehpeiensis as 3.5-4.0 (mean
in respects, particularly the antenna spine patterns 3.7) times longer than broad, thus slightly narrower and the indications of hairs or lack thereof on the than in the populations of M. ruttneri examined. medial extension of the leg 4 basipodite. Whether Otherwise there is no discernible difference from or not the populations from these mutually distant M. ruttneri. Subsequent descriptions of Asian localities are all conspecific, the description of each Mesocyclops populations ascribed to M. pehpeien-
differs in several respects from M. ruttneri. sis by Tai & Chen (1979) and by Kim & Chang
" In Sri Lankan "M. pehpeiensis (Dussart & (1989) unfortunately have not provided sufficient
Fernando, 1988) the caudal side of antenna article detail to clarify the problem (Reid & Kay, 1992). In
1 is shown as having a group of 5 tiny spines more view of the demonstratedpresence in Asia of sever-
less the level of the or at more proximal medialseta, al populations that differfromeach other in a series in contrast to the single or double row of 6-14 of microcharacters but that are all more or less con- small spines near the corresponding location and gruent with Hu's description, comparison of topo- the additional2-4 small spines near the distal cau- type material is essential for a final determination
dal that margin are always present and easy to see of the identity of his species with M. ruttneri, or in M. ruttneri. with subsequently described populations of Meso-
Figures by Dussart & Fernando (1986, locality cyclops from Asia. - 183 Bijdragen tot de Dierkunde, 63 (3) 1993
non-lacustrine in of Mesocyclops papuensis Van de Velde, 1987 from aquatic fauna of wetlands most
North America is known. Researchers Papua New Guinea is also very similar to M. rutt- inadequately
differences in the be alert for the of neri. There are slight antenna should possibility encountering
of the article 1 spine pattern, M. papuensis having a additional species genus.
of small the caudal side at diffuse group spines on the level of the medial setae (Van de Velde, 1987:
Genus char, emend. 159, figs. 45, 47) rather than the single or double Diacyclops Kiefer, 1927, 1928,
the Morton, 1985, Reid et 1989 rows of spines in M. ruttneri. In M. papuensis, al., medial terminal spine of leg 4 endopodite article 3
of its Diacyclops bernardi (Petkovski, 1986) has a few (4) large spinules near the midlength lateral margin, but in M. ruttneri nearly the entire (Figs. 6-8) length of the margin of this spine is spinulate. Also
Synonymy. — Mesocyclops bernardi Petkovski, 1986: 47, 48, in M. ruttneri, the leg 4 endopodite 3 medialtermi- 66-71, 72, Abb. 22-33; 1988: 40, 63; Sket, 1988: 79, table 1; nal spine is always distinctly longer than the lateral Reid, 1990: 180, table I; Marten, 1990b: 681. in terminal spine, but these spines are subequal Mesocyclops (?) bernardi; Dussart, 1987: 153, 155.
M. papuensis. Mesocyclops papuensis of course also resembles the general description of M. peh- Material. - USNM 251979, 1 9 on slide, Swale No. 1, Mail Box
274, Chef Menteur Highway, east New Orleans, Louisiana, peiensis. U.S.A., 14 November 1989, coll. G.G. Marten. USNM 259530, These records of M. ruttneri from the southern 1 Ç , ethanol-preserved, pond at side of Mérida-ProgresoHigh-
U.S.A. and are the first outside Austria. The Asia PEMEX 16 way, km 27, in front of plant, Mérida, Mexico, July Viet discoveries in China, Thailand, and Nam 1992, coll. M.F. Suárez and L. Bracamonte. probably represent native populations. The popula-
- of Louisiana tion in Austria was undoubtedly introduced into Description of female. Length
1.01 of Mérida 0.95 the greenhouse, and the species has never been col- specimen mm; specimen mm. lected elsewhere in Europe (Kiefer, 1981). The Habitus (Figs. 6a-c) as described by Petkovski populations in Louisiana and Mississippi were like- (1986), except pediger 4 laterally bilobed. Pediger 5
broad in dorsal view ly introduced, possibly either along with imported (Figs. 6a-c) approximately as
with of little tropical aquatic plants or fish, or somehow as broadest part genital segment, produced
collected ornamentation 2 dorsal rice. Wild populations of this species were posteriorly, lacking except in in ricefields Loui- hairs midline. Genital 1990 and 1991 near Jennings, near segment (Figs. 6a-d)
138 broad broadest with siana, about 300 km west of New Orleans, and near 125 /xm long, /¿m at part,
Cleveland, Mississippi, about 400 km north of New 3 dorsal papillae and 3 dorsal hairs, otherwise lack-
Orleans. The species seems to be well established ing ornamentation. Seminal receptacle (Fig. 6d), and probably spreading in the region. Adults origi- anterior outline difficult to distinguish, posterior
in both nally collected in New Orleans were introduced into expansion little developed specimens.
Jennings ricefields in 1990 and into both Jennings Copulatory pore (Figs. 6d, e) tiny, unsclerotized; and Cleveland ricefields in 1991 as part of mos- pore-canal slender, slightly curved; medial half of
quito-control experiments (G.G. Marten, personal margin of each lateral canal curved posteriorly, communications, 1990, 1991). lateral half slightly recurved anteriorly. The struc-
Mesocyclops ruttneri, like M. americanus Dus- tures of the pore and canals were not distinguished
Abb. but sketch of sart, 1985, is a member of the leuckarti-group. It is by Petkovski (1986: 67, 24), a
sent B.H. Dussart the second member of this group, after M. america- a paratype female by (in litt.,
shows structure similarto thoseof the nus, to be recorded in the contiguous U.S.A. and 1991) a speci-
and Mexico. rami Canada. Therefore the assumption by Dussart mens from Louisiana Caudal
times than broad, lack- (1985) and Dussart & Fernando (1990) that all (Figs. 6a, g, h) 2.65 longer records of M. leuckarti in these countriesmust refer ing ornamentation on medial surfaces and at base to M. americanus is not quite the case. The small of lateral seta, but having 6 tiny spines at base of 184 J. W. Reid New records of American Mesocyclops and Diacyclops bernardi
6. Fig. Diacyclops bernardi (Petkovski, 1986), from Louisiana, USNM 251979: a, habitus, dorsal; b, posterior prosome and anterior
and anterior urosome, right lateral; c, posterior prosome urosome, dorsal; d, genital segment, ventral; e, copulatory pore and pore-
ventral anal somite and caudal caudal lateral. Scale canal, lateral-oblique, showing curve; f, leg 5; g, rami,dorsal; h, right ramus, applies to Fig. 6a. Bijdragen tot de Dierkunde, 63 (3) - 1993 185
base of teeth of incisiva. Maxillule lateralmost terminal seta. In Louisiana specimen, spines near pars
right caudal ramus also with dorsoventral lateral (Fig. 7f) lacking spines on surface of palp. Claw of
row of 3 tiny spines and 1 papilla midway between maxilla (Fig. 7g) with stout teeth along middle of
otherwise similar base of ramus and lateral seta; left caudal ramus margin, to maxilla of D. navus.
with similar papilla but lacking spines; Mexican Maxilliped (Fig. 7h) with 1 longitudinal and 1 trans-
specimen with reverse arrangement. Specimen from verse row of spines on article 2.
Louisiana: both medial terminal caudal setae of Legs 1-4 (Figs. 8a-d) with triarticulate rami,
right caudal ramus broken; lengths of setae of left spine formula 2,3,3,3, seta formula 4,4,4,4/3. Leg
caudal in dorsal lateral medial- 1 much described Petkovski 67, Abb. ramus /¿m, 65, 36, as by (1986:
most to lateralmost terminal 82, 400, 312, 115. 27), additionally with medial expansion of basi-
Specimen from Mexico: lengths of caudal setae, podite haired, 2 short diagonal rows of spines on
dorsal 58, lateral 35, medialmostto lateralmost ter- anteriorsurface of basipodite, and lateral spines of
minal 73, 380, 296, 104 ¿im. Lateralmost terminal exopodite somewhat stouter than in specimens
caudal seta about 2 times thicker at base than me- from San Andrés. Legs 2 and 3 similar to each
dialmost terminal caudal seta. All setae with uni- other, strikingly larger than legs 1 and 4, exopodites
setules. from and lateral of formly fine, closely set Specimen spines exopodites very stout. Leg 4
Louisiana with paired egg sacs, each with 17 eggs. much as described by Petkovski (1986), except
Antennule(Figs. 6a, 7a, b) of 16 articles in both lateral spines of exopodite of specimens from Loui-
specimens, articles numbered in Roman numerals siana and Mexico relatively stouter. Coxa-basi-
(with numbers of setae in parentheses): 1(8), 11(4), podite of Louisiana specimen ornamented as illus-
111(2), IV(5), V(3), VI(1), VII(2), VIII(l), IX(1), trated. Leg 4 basipodite of specimen from Mexico
X(0), XI(1), XII(l), XIII(l), XIV(2), XV(1), with similar ornamentation, except row of spines
border XVI(7). Article 6 with spine, articles 12 and 15 each on posterodistal longer, extending along with aesthetasc; slender aesthetasc of article 15 middle Vi of border. Louisianaspecimen: right leg reaching nearly midlength of article 16. Hyaline 4 exopodite article 3 with 2 setae and 1 socket; left membrane of article 15 narrow, finely serrate; exopodite article 3 with 4 setae, next distalmost seta membrane of article both articles with 16 broad, more coarsely ser- short; additionally 3 large spines. rate distally. Surface of articles otherwise lacking Specimen from Mexico with 4 setae on each exopo- ornamentation, except row of small spines on arti- dite article 3. cle 1. Numberof 16articles resulting from apparent Leg 5 (Fig. 6f), lateral setae of article 1 of both fusion of former articles 13 and 14, by analogy with legs broken in Louisianaspecimen and, in specimen location and numbers of setae of antennules of from Mexico, slightly longer than terminal seta of
Mesocyclops leuckarti as redescribed by Van de article 2. Inboth specimens, terminal spine and seta
Velde and of rede- of article about of (1984a) Diacyclops navus as 2 reaching midlength genital seg- scribed by Reid et al. (1989). ment. Leg 6 (Fig. 6b) consisting of ovoid plate lack-
Antenna (Figs. 7c, d) similar in general structure ing surface ornamentationand bearing 2 spines and and number of setae to antenna of D. navus (Reid 1 short dorsal seta. et al., 1989: 340, fig. 2b); article 1 bearing, in addi- tion to 2 short slender setae and 1 long stout seta on Discussion. - Examinationof the specimens of D.
border: distal 1 short row of spines along proximal bernardi from Louisiana and Mexico confirms and part of lateral margin and 2 rows of spines along reinforces Petkovski's (1986) observation that this proximal part of medial margin, 1 short longitudi- species combines morphological features of several nal row of spines on frontal side near lateral margin genera. These generaare Diacyclops, Mesocyclops, and 2 curved of caudal longitudinal rows spines on and Thermocyclops as presently defined; cf. recent side. diagnoses of Diacyclops by Kiefer (1978) and
Mandible (Fig. 7e) similar to that of D. navus, Morton (1985), and of Mesocyclops and Thermocy-
with of slender lacking spines near palp, 1 row clops by Kiefer (1978). Petkovski (1986) assigned 186 J. W. Reid - New records of American Mesocyclops and Diacyclops bernardi
Fig. 7. Diacyclops bernardi (Petkovski, 1986), from Louisiana,USNM 251979: a, antennule; b, antennule articles 15-16; c, antenna
article article 1, frontal side; d, antenna 1, caudal side; e, mandible; f, maxillule; g, maxilla; h, maxilliped. Bijdragen tot de Dierkunde, 63 (3) - 1993 187
Fig. 1 8. Diacyclops bernardi (Petkovski, 1986), from Louisiana, USNM 251979: a, left leg and coupler, anterior; b, left leg 2 and
4 part of coupler, anterior; c, left leg and coupler, posterior; d, right leg 4 exopodite article 3, posterior. Plumage of most setae omitted for clarity. 188 J. W. Reid New records of American Mesocyclops and Diacyclops bernardi
the species to Mesocyclops on thebasis of the struc- case in Mesocyclops. On the frontal side there is a
and the short tures of the seminal receptacle leg 4 coupler, row of 8 spines at about the proximal third,
although he called attention to the similarity of the rather than the long longitudinal row of some 20 or
terminal spines of the leg 4 endopodite and the ter- more spines usually present in Mesocyclops, and
minal placement of the medial spine of leg 5 to spe- the proximal transverse row of tiny spines found in
is The cies of Thermocyclops. Dussart (1987), having ex- Mesocyclops not present in D. bernardi. pat-
amined some of Petkovski's specimens, advanced a tern in D. bernardiis less ornate than in many spe-
tentative opinion that the species is better placed in cies of Thermocyclops, although all the spines are
Thermocyclops. Dussart mentioned that the struc- long, rather than being reduced to tiny "teeth" as
of the seminal and the fine in of ture receptacle leg 5, ser- many species the latter genus (Fiers & Van de
ration of the membraneof the The of bernardi how- hyaline antennule, Velde, 1984). pattern D. is,
and the ornamentationof article 1 of the antenna ever, very similar to the patterns of Diacyclops bi-
agree better with Kiefer's (1927, 1978) diagnoses of cuspidatus (Claus, 1857) sensu stricto and D. bicus-
that genus. pidatus odessanus (Schmankevitch, 1875), as pre-
The distinctly sclerotized lateral arms and the sented by Fiers & Van de Velde (1984), and also to
shape of the posterior expansion of the seminal re- D. navus as redescribed by Reid et al. (1989), and
resemble of all three D. ceptacle some species genera, palustris.
e.g., Mesocyclops meridianus (Kiefer, 1926), M. The terminal article of the antennuleof D. ber-
reidae, Thermocyclops tenuis(Marsh, 1910), T. in- nardi is about 4 times longer than broad, similar to
versus Kiefer, 1936b, Diacyclops palustris Reid, the usual proportions of this article in species of
and D. The of the 1988, navus. structure receptacle Mesocyclops. The corresponding article in Ther-
therefore supports no argument for inclusion of mocyclops is usually about 2 times longer than
this species in a particular genus. broad, serving as a useful distinction between the
The antennule is composed of 17 articles in spe- two genera (J.F. Saunders, III, pers. comm., 1984;
of cies Mesocyclops and Thermocyclops, and I am Reid, 1985). In Diacyclops, the proportions of this .
unaware of of reduction of article article differ from 3.2 times any example considerably, about
number in those genera. Mostof the specimens col- longer than broad in D. navus and D. palustris to lected from San Andrés have theantennuleof 16 ar- as short as 2 times longer than broad in D. lan- ticles, with one having 17 articles (T.K. Petkovski, guidoides (Lilljeborg, 1901) sensu lato as rede- in The of litt., 1991). number 16articles appears to scribed by Reid et al. (1991). be most characteristic for D. bernardi. Reduction Inknown species of Mesocyclops and Thermocy- and variationin antennulearticle numberis a com- clops, the medialmost terminal caudal seta usually mon phenomenon in species of Diacyclops. is much longer than the caudal rami and is con-
The structure of the hyaline membraneof the an- sistently longer than the lateralmost terminal cau- tennule likewise offers little basis for generic assig- dal seta. In species of Diacyclops this seta is usually nation because three all genera include some species short, as in D. bernardi.
unnotched with a similar finely serrate, membrane. Many species of Diacyclops possess a dorsoven-
The membrane is broader than in known species of tral row or comb of small spines on the lateral sur-
Thermocyclops and Diacyclops. face of the caudal ramus about equidistant between
The ornamentationof article 1 of the antenna of the base of the ramus and the lateral seta, similar to
D. bernardidiffers of of from substantially from the basic pat- the spines the specimens D. bernardi tern found in Mesocyclops (Van de Velde, 1984a, Louisianaand Mexico. Such arow of spines has not b). On the proximal part of the medial margin, to my knowledge been described in species of there is a row of large spines rather than the tiny Mesocyclops or Thermocyclops. However, I have
found at this location in of at this location in spines many (though not observed a short row spines all) species of Mesocyclops, and these spines do not two of hundreds of specimens of M. ruttneri ex- continue as a row on the caudal side as is the usual amined from Louisiana and Mississippi. Bijdragen tot de Dierkunde, 63 (3) - 1993 189
The leg 4 endopodite article 3 in Mesocyclops is Acknowledgements
usually 3-4 times longer than broad, and the two
terminal spines are subequal in length. The cor- Several colleagues provided cordial assistance, answering in-
Drs. responding article in D. bernardi is short by com- quiries and lending or donating specimens: Bernard H. Dussart, C. Herbert Fernando, Teruo Ishida, Gerald G. Marten, parison and the terminal spines are of greatly dif- Trajan K. Petkovski, James R. Reddell, Marco F. Suárez, Vu in of ferent lengths, as is the case many species Sinh Nam, and Harry C. Yeatman. Dr. Hans-Walter Mittmann and Thermocyclops Diacyclops. Dr. lent specimens and some of Friedrich Kiefer's original of the The large triangular spiniform processes sketches from the Kiefer Copepod Collection in the Staatliches
leg 4 coupler of D. bernardi are similar to structures Museum fiir Naturkunde Karlsruhe. Dr. Gerald G. Marten and
of and differ- his technical assistants made the extensive collections which present in many species Mesocyclops
provided the impetus for this report. ent from the roundedprocesses in many Thermocy-
clops. Species of Diacyclops usually have no or only
smallrounded processes on the leg 4 coupler, which References may be ornamented also with small spines on its
surface. Burmeister, H., 1834. Beitràgezur Naturgeschichte der Ranken- The robustness of the swimming legs and their fiisser (Cirripedia): 1-60, Taf. 1, 2 (G. Reimer, Berlin). lateral spines, especially legs 2 and 3 in D. bernardi, Claus, C., 1857. Das genus Cyclops und seine einheimischen
is an unusual feature for all these and genera Arten: 1-40, Taf. I—III (Inaugural-Dissertation, Marburg).
United may be an adaptation for a benthic and possibly Dana, J.D., 1853. Crustacea. In: States ExploringExpe-
dition the under primarily subterranean existence. during years 1838, 1839, 1840, 1841, 1842,
the command of Charles 687-1618 of Wilkes, U.S.N., 13(2): In the present state of our concept of genera (C. Sherman, Philadelphia). Folio Atlas, pis. 1-96, pub- Cyclopidae, the structure of the fifth leg is still af- lished 1855. forded bernardithe medial great importance. In D. Dussart, B.H., 1985. Le genre Mesocyclops (Crustacé, Copé- of article 2 of the fifth is subtermi- spine leg clearly pode) en Amérique du Nord. Can. J. Zool., 63(4): 961-964.
nal in accordance with the diagnoses of Thermocy- Dussart, B.H., 1987. Sur quelques Mesocyclops (Crustacea,
Copepoda) d'Amérique du Sud. Amazoniana, 10(2): 149- clops and Diacyclops, but not with Mesocyclops in 161. which the medial spine is inserted about midlength Dussart, B.H. & C.H. Fernando, 1985. Les Copépodes en Sri of the article. The terminal spines of the fifth legs Lanka (Calanoïdes et Cyclopoïdes). Hydrobiologia, 127(3): of several included in species presently Diacyclops 229-252.
are comparatively long. For instance, the spine in Dussart, B.H. &C.H. Fernando, 1986. The Mesocyclops species
58: D. navus frequently approaches or exceeds the problem today. Syllogeus, 288-293.
Sur of terminal and reach Dussart, B.H. & C.H. Fernando, 1988. quelques Mesocy- length the seta may midlength clops (Crustacea, Copepoda). Hydrobiologia, 157(3): 241 — of the genital segment in the female and nearly to 264. leg 6 in the male (Reid et al., 1989). Dussart, B.H. & C.H. Fernando, 1990. A review of the taxono-
In the nature of several characters summary, of five Ontario of freshwater my genera cyclopoid Copepoda transfer of this has persuaded me to propose spe- (Crustacea). Can. J. Zool., 68(12): 2594-2604.
Fiers, F. & I. van de Velde, 1984. Morphology of the antenna cies to the genus Diacyclops. These are: the number and its importance in the systematics of the Cyclopidae. of articles of the antennule, which is usually 16 Crustaceana, Suppl. 7(2): 182-199. rather than 17; the ornamentationof the antenna; Forbes, S.A., 1891., On some Lake Superior Entomostraca. the structure of the seminal receptacle; the dor- Annual Report of the Commissioner of Fish and Fisheries lateral surface soventralrow of small spines on the for 1887: 701-718, pis. I—IV (Government Printing Office, of the caudal ramus; and the structures of the Washington).
fourth and fifth legs. Gurney, R., 1933. British freshwater Copepoda, 3: 1-334 (Ray Society, London). The record of D. bernardi from New Orleans, Herrick, C.L., 1887. Contribution to the fauna of the Gulf of from an intermittently floodedswale, was previous- Mexico and the South. Mem. Denison scient. Ass., 1(1): 3— ly reported by Marten (1990b). On San Andrés 56, pis. I—VII, IX.
collections were made from and a Island, pools Hu, Y.T., 1943. Notes on fresh-water copepods from Pehpei, cave (Petkovski, 1986). Szechwan. Sinensia, Nanking, 14(1-6): 115—128. 190 J. W. Reid - New records of American Mesocyclops and Diacyclops bernardi
9. Neue Acta Mus. maced. Sci. 19 Kiefer, F., 1926. Beitrâge zur Copepodenkunde(IV). (Crustacea, Copepoda). nat.,
Cyclops-Arten. Zool. Anz., 69(1/2): 21-25. (2/155): 39-64.
Kiefer, F., 1927. Versuch eines Systems der Cyclopiden. Zool. Reddell, J.R., 1965. A checklist of the cave fauna of Texas, I.
Anz., 73(11/12): 302-308. The Invertebrata (exclusive of Insecta). Texas J. Sci., 17(2):
Kiefer, F., 1928. Über Morphologie und Systematik der Siiss- 143-187.
wasser-Cyclopiden. Zool. Jahrb., Abt. Syst. Ökol. Geogr. Reddell, J.R., 1981. A review of the cavernicole fauna of
Tiere, 54(5/6): 495-556. Mexico, Guatemala, and Belize. Bull. Texas Meml. Mus., 27:
Kiefer, F., 1936a. Brasilianische Ruderfusskrebse (Crustacea 1-327.
J.R. & R.W. A Copepoda), gesammelt von Herrn Dr. Otto Schubart. V. Mit- Reddell, Mitchell, 1969. checklist and annotated
teilung. Zool. Anz., 116(1/2): 31—35. bibliographyof the subterranean aquatic fauna ofTexas. In-
Kiefer, F., 1936b. Brasilianische Ruderfusskrebse (Crustacea ternational Center for Arid and Semi-Arid Land Studies,
Dr. Copepoda), gesammeltvon Herrn Otto Schubart. II. Mit- Texas Technological College Water Resources Center,
teilung. Zool. Anz., 114(5/6): 129-133. Lubbock, Texas, Spec. Rep., 24: 1-48.
Kiefer, F., 1978. Das Zooplankton der Binnengewâsser, 2. Reid, J.W., 1985. Chave de identificaçâo e lista de referências
Freilebende Copepoda. Binnengewâsser, 26(2): 1-343. bibliográficas para as espécies continentais sulamericanas de
Kiefer, F., 1981. Beitrag zur Kenntnis von Morphologie, Tax- vida livre da ordem Cyclopoida (Crustacea, Copepoda).
onomie und geographischer Verbreitung von Mesocyclops Bolm. Zool., Univ. Sào Paulo, 9: 17-143.
leuckarti auctorum. Arch. Hydrobiol., Suppl 62 (Monogr. Reid, J.W., 1988. Copepoda (Crustacea) from a seasonally
Beitr.), 1: 148-190. flooded marsh in Rock Creek Stream Valley Park, Maryland.
Kim, H.S. & C.Y. Chang, 1989. Freshwater cyclopoid copepods Proc. biol. Soc. Wash., 101(1): 31-38.
(Cyclopoida, Cyclopidae) of Korea. Korean J. syst. Zool., Reid, J.W., 1990. Continental and coastal free-livingCopepoda
5(2): 225-256. (Crustacea) of Mexico, Central America and the Caribbean
Lilljeborg, W., 1901. Synopsis specierum hue usque in Suecia Region. In: D. Navarro L. & J.G. Robinson (eds.), Diver-
observatarum generis Cyclopis, sive Bidrag till en Öfversigt af sidad Biológica en la Reserva de la Biosfera de Sian Ka'an,
de inom Sverige iakttagna arterna af slàktet Cyclops. K. Quintana Roo, Mexico: 175-213 (Centro de Investigaciones
svenska VetenskAkad. Handl., (N. F.) 35(4): 1-118, pis. de Quintana Roo (CIQRO) & Program of Studies in Tropical
I-VI, 2 tables. Conservation, University of Florida; Chetumal, Quintana
Lim, R.P. &C.H. Fernando, 1985. A review of Malaysianfresh- Roo).
water Copepodawith notes onnew records and little known Reíd, J.W., 1992. Copepoda (Crustacea) from fresh waters of
species. Hydrobiologia, 128(1): 71-89. the Florida Everglades, U.S.A., with a description of Eucy-
Marsh, C.D., 1910. A revision of the North American species of clops conrowae n. sp. Trans, amer, microsc. Soc., 111(3):
Cyclops. Trans. Wisconsin Acad. Sci. Arts Lett., 16(2): 229-254.
1067-1136. Reid, J.W., S.G.F. Hare & R.S. Nasci, 1989. Diacyclops navus
Marten,G.G., 1989. A survey of cyclopoidcopepods for control (Crustacea: Copepoda) redescribed from Louisiana, U.S.A.
of Aedes albopictus larvae. Bull. Soc. Vector Ecol., 14(2): Trans, amer, microsc. Soc., 108(4): 332-344.
& 232-236. Reid, J.W. B.H. Kay, 1992. Mesocyclops guangxiensis, new
Marten, G.G., 1990a. Issues in the development of cyclops for species, and new records of four congeners (Crustacea:
mosquitocontrol. In: M. F. Uren, J. Blok & L. H. Mander- Copepoda: Cyclopidae) from China, Laos, and Viet Nam.
son (eds.), Proceedings of the 5th Symposium on Arbovirus Proc. biol. Soc. Wash., 105(2): 331-342.
Research in Australia: 159-164 (CSIRO & Queensland Inst. Reid, J.W., E.B. Reed, J.V. Ward, N.J. Voelz & J.A. Stanford,
1. Med. Research, Queensland). 1991. Diacyclops languidoides (Lilljeborg, 1901) s. and
Evaluation of Ae- Marten,G.G., 1990b. cyclopoid copepods for Acanthocyclops montana, new species (Copepoda, Cyclopoi-
des albopictus control in tires. J. amer. Mosquito Control da), from groundwater in Montana, U.S.A. Hydrobiologia,
Ass., 6(4): 681-688. 218(2): 133-149.
Morton, D.W., 1985. Revision of the Australian Cyclopidae Sars, G.O., 1914. An account of the Crustacea of Norway with
(Copepoda: Cyclopoida), I. Acanthocyclops Kiefer, Diacy- short descriptions and figures of all the species. VI. Copepoda
V clops Kiefer and Australocyclops, gen. nov. Aust. J. mar. Cyclopoida. Parts and VI. Cyclopidae (continued): 57-80
Freshwat. Res., 36: 615—634. (Bergen Museum, Bergen).
Pesce, G.L., 1985. Amsterdam Expeditions to the West Indian Schmankevitch, V.I., 1875. Quelques Crustacés des eauxsalées
Islands, Report 45. Cyclopids (Crustacea, Copepoda) from et douces et leurs rapports avec l'élément environnant. Mém.
West Indian habitats. groundwater Bijdr. Dierk., 55(2): 295- Soc. Naturalistes nouvelle Russie, Odessa (= Zap. novoross.
323. Obshch. Estest.), 3(2): 1-391, pis. 1-5.
Petkovski, T.K., 1986. Zur Taxonomie des Genus Mesocyclops Sket, B., 1988. Fauna ofthe hypogean waters ofthe San Andres
G.O. Sars 1914 (Crustacea, Copepoda Cyclopoida) in der (Saint Andrews) Island, Colombia. Biol. Vestn., 36(2): 77—
Neotropis. Acta Mus. maced. Sci. nat., 18(3/149): 47-49. 82.
Petkovski, T.K., 1988. Zur Cyclopidenfauna Kolumbiens Smith, K.E. & C.H. Fernando, 1978. The freshwater calanoid Bijdragen tot de Dierkunde, 63 (3) - 1993 191
and cyclopoid Crustacea of Cuba. Can. J. Zool., 56(9): characters in Mesocyclops, with African species as an ex-
2015-2023. ample. Crustaceana, Suppl. 7(2): 404-419.
Smith,K.E. & C.H. Fernando, 1980. Guía para los copépodos Velde, I. van de, 1987. New Mesocyclops species (Copepoda,
de 1 de New (Calanoida y Cyclopoida) Cuba: -28 (Academia Cien- Cyclopidae) from Papua Guinea. Bull. Inst. roy. Sci.
cias de Cuba, Havana). nat. Belg., (Biol.) 57: 149-162.
Tai, A.-y. & G.-x. Chen, 1979. Cyclopoida. In: C.-j. Shen (ed.), Yeatman, H.C., 1977. Mesocyclops ellipticus Kiefer from a
Fauna Sinica: Crustacea: Freshwater Copepoda: 301-420 Mexican cave. In: J.R. Reddell (ed.), Studies on the caves and
(Science Press, Peiping). cave fauna of the Yucatan Peninsula. Bull. Ass. mex. Cave
6: Velde, I. van de, 1984a. Revision of the African species of the Stud., 5-7.
genus Mesocyclops Sars, 1914 (Copepoda: Cyclopidae).
Hydrobiologia, 109(1): 3-66.
Velde, I. van de, 1984b. Introduction of new identification Received: 26 February 1993