BELKI, A NEW FROM COZUMEL. MEXICO (CONCHOSTRACA: ). WITH A REVIEW OF CENTRAL AND SOUTH AMERICAN SPECIES OF THE GENUS EULIMNADIA

INVERT, ZOOLOGY SECTION LIBRARY UCM

JOEL W. MARTIN

Made in United States of America Reprinted from JOURNAL OF BIOLOGY Vol. 9, No. 1, Febmary 1989 Copyright ft ] 989 bv the Crustacean Society JOURNAL OF CRUSTACEAN BIOLOGY. 9(1): 104-114. 1989

EULIMNADIA BELKI, A NEW CLAM SHRIMP FROM COZUMEL, MEXICO (CONCHOSTRACA: LIMNADIIDAE), WITH A REVIEW OF CENTRAL AND SOUTH AMERICAN SPECIES OF THE GENUS EULIMNADIA

Joel W. Martin

ABSTRACT A new species of Eutimnadia, E. belki, is described from the Chankanaab National Park in Cozumel, State of Quintana Roo, Yucatan Peninsula, Mexico. The species also occurs in Ven­ ezuela. Eulimnadia belki can be distinguished from all other species of the Limnadiidae in the New World by the unusual morphology of the eggs. The morphology of the external egg shell, a promising taxonomic character, is compared to that of all species oi Eulimnadia known from Central and South America.

The conchostracan family Limnadiidae is (male holotype, LACM No. 85-188.1; female allotype, known from nearly all comers of the world. LACM No. 85-188.2; 12 female and 1 male paratypes. LACM No. 85-188.3). Members of the family are easily recognized Specimens of other Eulimnadia were borrowed from by the presence of a pedunculate frontal or­ various museums and from private collections as fol­ gan on the midline of the head region (with lows. British Museum (Natural History): E. chacoensis few exceptions), the absence of a supportive Gurney, syntypes, 1928.2.23.1-15, Makthlawaiya, ridge (fornix) on either side of the rostrum, Paraguayan Chaco, collected by G. S. Carter; E. an- tillamm (Baird), 52-23, Santo Domingo, collected by and the presence of few to many concentric M. Salle. Hungarian Museum of Natural History, Col- growth lines on the laterally compressed lectio Dadayana PhvUopoda: E. amillarum (Baird), valves. The genus Eulimnadia, containing Santo Domingo, CDP l/C-133, 1913.-11; E. brasil- about 40 described species, is the most di­ iensis Sars, Iparanga, CDP I/C-134,1913.-116; £. geayi Daday, Venezuela, CDP I/C-143, 1913.-125; £. "co- verse genus not only of the Limnadiidae but lumbica" (named in vial only, see Forro and Brtek, possibly of the entire order Conchostraca 1984; = E. geayi; see Discussion), CDP I/C-136, 1913.- (see Belk, 1982; Straskraba, 1965) (=order 118. Private collections of Dr. Denton Belk: Eulim­ Spinicaudata; see Fryer, 1987). In Septem­ nadia sp. A [DB 305], Argentina, Catamarca, Rt. 1, ber of 1985, a small lot of undescribed lim- km 45, 30 December 1973, collected by A. Hulse; Eu­ limnadia sp. B [DB 632], Paraguay, Chaco Parque Na- nadiid conchostracans belonging to the gQ- cional Defensoresdel Chaco, Tribo Nuevo, "econtrado mxs Eulimnadia was collected from a shallow en regiones bajas de laguna recien inundada," 23 No­ surface pool in the Parque de Chankanaab, vember 1984, collected by T. Bonace and D. Drenner. Quintana Roo, Cozumel, Mexico; the lot was later brought to my attention by Jill Eulimnadia belki. new species Yager. The species is described below. Figs. 1-3, 4A, B Material. —Mexico: Parque de Chankanaab, Cozumel, MATERIALS AND METHODS State of Quintana Roo; 3 33, 15 99, collected by D. Williams, J. Bozanic, 19 September 1985. Conchostracans (15 females, 3 males) were collected by D. Williams and J. Bozanic from a shallow surface Measurements.—Holotype 3 3.75 mm length, 2.38 mm pool in the Chankanaab National Park, Cozumel, Mex­ height; other 3S both 3.63 mm length, 2.19 mm height ico. Approximate area of the pool was 10 x 4 m with (I specimen destroyed for SEM). Allotype 9 4.94 mm a depth of 16 cm. Salinity of the pool at the time of length, 3.19 mm height; other 95 3.56-4.56 mm length, collection was less than 1 ppt. Conchostracans were 2.19-2.75 mm height (2 specimens destroyed for SEM). initially ixed in fresh-water Formalin and later trans­ ferred to 70% ethanol. Carapace.—Carapme (Figs. lA, B, D, 3A) Illustrations were prepared using a Wild M-5A ste­ in both sexes oval, with 4 concentric growth reoscope and M-11 compound microscope, both with lines in males and 4 or 5 in females. Umbo camera lucida. Specimens were prepared for scanning lacking. Area between growth lines minute­ electron microscopy (SEM) following procedures out- ly granulate except for narrow band on either Uned by Felgenhauer (1987) but without postfixation in osmium and with 100% ethanol used as the tran­ side of growth lines (Fig. 3 A). Posterior half sitional fluid. All specimens have been deposited in of dorsal hinge more or less flatan d straight. the Natural History Museum of Los Angeles County Anterior carapace narrow in dorsal view,

104 MARTIN: NEW EULIMNADIA FROM COZUMEL, MEXICO 105

Fig. 1. Eulimnadia belki, new species, holotype male (A-C, F) and allotype female (D, E). A, male, lateral view; B, male, dorsal view; C, male, head region, lateral view; D, female, lateral view, with eggs slightly enlarged in figurea t upper right; E, female, head region, lateral view; F, male, caudal region, lateral view. Abbreviations: am = adductor muscle, an 1 = first antenna (antennule), an2 = second antenna, ce = compound eye, cf = caudal furca, fo = frontal organ, hp = hepatopancreas, la = labrum, oc = ocellus, p = approximate position of frontal pore, tf = telsonal filaments. Scale bar = 0.5 mm for A-F.

widening rapidly posteriorly (Fig. IB). Max­ Head Region.—Male head region (Fig. IC) illary gland (mg) elongate and surrounding with large frontal organ (fo) arising from adductor muscle (am). Females slightly protuberance just posterior to compound eye larger and more inflated, especially so if (ce); area dorsal to compound eye with large ovigerous. "empty" space, possibly connecting to out- 106 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 1, 1989

Fig. 2. Eulimnadia belki, first thoracopods (claspers) of holotype male. A, left thoracopod 1, anterior surface; B, right thoracopod 2, anterior surface. Scale bar = 0.5 mm. Setal types and suckerlike distal prominence not drawn to scale. side environment via pore (p) located on bearing internally ocellus; hepatopancreas midline just dorsal and anterior to com­ as in male. pound eyes, head region being wide in fron­ tal view (Figs. IC, 3B). Small low rounded Antennae. —First antenna (antennule) (anl) lobe just dorsal to compound eye. Rostrum extending to about fourth segment of pos­ produced into long narrow blunt projection terior antennal flagellum, indistinctly seg­ bearing internally naupliar eye or ocellus mented with 8 or 9 lobes bearing short sen­ (oc) variable in shape. Hepatopancreas (hp) sory setae. Second antenna (an2) with extending slightly into rostrum and into indistinct segmentation of peduncle and with dorsal (proximal) part of labrum (la). Fe­ 9 segments on both anterior and posterior male head region (Fig. IE) similar, with flagella, each bearing short, sharp dorsal "empty" area and midfrontal pore, but with spines and longer natatory ventral setae rostrum not as narrow or as produced as in (Fig. IC, E). male, terminating in obtuse blunt angle Male Thoracopods.—First thoracopod (Figs. MARTIN: NEW EULIMNADIA FROM COZUMEL, MEXICO 107

w _^ " .^^

1, ,' ^^^ • •• - / / -^ .ST*- . tf ^

^^

^^^-jv^ F N\ V -^?^

Fig. 3. Scanning electron micrographs of selected characters of Eulimnadia belki. A, anteroventral part of left valve of male, showing 4 concentric growth lines and details of sculpturing, x 120; B, frontal view of male rostrum, x 150; C, first male thoracopod (clasper), posterior surface, x 300; D, second male thoracopod (clasper), X 190; E, tip of clasper fingers showing suckerlike prominence (indicated by unlabeled arrow in D) and stout, flat-tipped spines and serrate spines (far right), x 1,300; F, caudal region of female, x 130; G, high magnification of posterior spines indicated by unlabeled arrow in F, x 1,500; H, distal region of left ramus of caudal furca (note small dorsal spinules), x 430. Abbreviations: an = second antenna; cf = caudal furca; p = frontal pore; r = rostrum; tf = telsonal filaments.

2A, 3C) stout, with obvious indentation in sodistal projection and stout, flat-tipped "hand" of clasper proximal to area bearing spines on clasping border; immovable fin­ stout teeth. Movable finger of clasper ger with larger stout flat-tipped spines op­ smoothly curving with small suckerlike dor- posite those of movable finger and with long 108 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. I, 1989 serrate spines proximal to stout ones (Figs. Venezuelan population taken by Guido Pe- 2A, 3C). Endite 4 (see Fryer, 1987, for ter­ reira). minology of clasper endites in nonlynceid Etymology. —Named in honor of Dr. Den­ conchostracans) short with simple setae; en­ ton Belk for his many contributions to the dite 5 (palp) 2-segmented, each segment biology of the and for his kind bearing terminal setae, with total length and unabated assistance offered to students (both segments) slightly exceeding length of and colleagues over the course of his career. movable finger. Second thoracopod (Figs. 2B, 3D) similar in general shape and spi- DISCUSSION nation, but with 2-segmented palp much The of the family Limnadi- longer than in first thoracopod, combined idae, and especially of Eulimnadia, is con­ length of segments nearly twice length of fused and badly in need of revision. Pre­ movable finger. Spination of tip of clasper vious descriptions of members of the family, fingers similar to first thoracopod (Fig. 3E). with few exceptions such as the elegant work Telson. —Posterior region of trunk (see of G. O. Sars (1896), are lacking in detailed Bowman, 1971; Schminke, 1976, for con­ morphology of characters that authors have trasting opinions on terminology) with 12- previously used to distinguish species. Thus, 14 large sharp spines, each minutely serrate it is often difficult to assign a specimen to under high magnification (Figs. IF, 3F, G), a known species, and the erection of new terminal spine of series much larger than species necessitates examination of mu­ preceding ones. Caudal furca (cf) well de­ seum holdings. Compared to the number of veloped, with 10-12 plumose setae and nu­ species of Eulimnadia known from North merous small spinules on distal upturned America (see Pennak, 1978; Belk, 1989), fourth of furca (Figs. IF, 3F, H). Telsonal relatively few species of Eulimnadia have filaments (tf) delicate, plumose only on dis­ been described from Central and South tal one-half to two-thirds, arising from about America (see Table 1). Loftier (1977, 1981) level of third posterior spine (Figs. IF, 3F); listed only Eulimnadia brasiliensis Sars from posteroventral maiigln of trunk (below in­ tropical and southern South America, re­ sertion of caudal furcae) with well-devel­ spectively, and E. antillarum (Baird) from oped blunt spine (characteristic of genus). the Caribbean; E. antillarum is also known from Mexico (Daday, 1926; Hartland-Rowe, Eggs.—StonX, approximately cylindrical, 1982) and southern Brazil (Lilljeborg, 1889). with inflated rims and with smoothly In addition to these two species, E. cha- rounded ridges defining narrow valleys ex­ coensis was described by Gumey (1931) tending parallel to each other along length from Paraguay, E. geayi was described by of cylinder from rim to rim (Figs. 1D, 4A, Daday (1926) from Colombia, Venezuela, B). Ends slightly concave, with parallel and Mexico, and E. texana has been re­ grooves and ridges similar to those on cyl­ ported from Mexico (Moore, 1965) and Sao inder, not radiating outward from center but Paulo, Brazil (Daday, 1926; Lutz, 1929). An running approximately 90° to axis of cyl­ undescribed species of Eulimnadia, collect­ inder and with groove occasionally extend­ ed by Daday in "Columbia" [sic = Colom­ ing through rim and merging with longitu­ bia] and bearing the label E. columbica, is dinal groove of cylinder; one end usually housed in the Hungarian Museum of Nat­ with single groove running obliquely to oth­ ural History (see Forro and Brtek, 1984). er end grooves (Fig. 4B). Rims of cylinder Finally, an undescribed species of Eulim­ flaring out slightly; egg diameter conse­ nadia was reported from Venezuela by Mar- quently somewhat wider at ends than if galef (1961) and several additional un­ measured at midpoint. Egg surface textured named species exist in the private collections with numerous minute pores. of Dr. E>enton Belk, Our Lady of the Lake Type Locality.—¥mque de Chankanaab, University, San Antonio, Texas. Quintana Roo, Cozumel, Mexico, shallow Compared to the above species, E. belki surface pool. is somewhat unusual in possessing an acute­ Range.—Type locality and Venezuela (based ly produced male rostrum and a very long on scanning electron micrographs of eggs of palp on the second clasper. In Eulimnadia MARTIN: NEW EULIMNADIA FROM COZUMEL, MEXICO 109

Fig. 4. Egg morphology oi Eulimnadia belki and other species oi Eulimnadia. A, two eggs oiE. belki showing longitudinal grooves along cylindrical axis, x 300; B, eggs of £•. belki with end pieces showing, x 350; C, spherical egg oiE. brasiliensis, x 500; D, stout cylindrical eggs oiE. geayi showing pentagonal appearance, x 350; E, egg oiE. geayi (note one raised and one flattened end), x 500; F, egg of £^. geayi showing detail of raised end, x 500. sp. (Margalef, 1961: fig.6A , B) the proximal er illustrated by Daday (1926); that species segment of the palp is shown much shorter differs from E. belki in having only 2 growth than in E. belki, and the male rostrum is lines on the valves and in having the pos- short and blunt rather than acutely pro­ terodorsal caudal region nearly devoid of duced. Eulimnadia antillarum was only spines (Fig. 5A). The body of Eulimnadia partially illustrated by Baird (1852) and lat­ antillarum is also slightly larger relative to 110 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 1, 1989 MARTIN: NEW EVUMNADU FROM COZUMEL, MEXICO 111

Table 1. Spedes oi Eulimnadia reported from Central or South America and the Caribbean.*

Species Locality Eulimnadia antillarum (Baird, 1852) Santo Domingo, West Indies; Mexico; Bra/il Eulimnadia belki, new species Cozumel, Mexico; Venezuela Eulimnadia brasiliensis Sars, 1902 Brazil Eulimnadia chacoensis Gumey, 1931 Paraguay Eulimnadia "columbica" (ms name only) Colombia (see Forro and Brtek, 1984) Eulimnadia geayiDaday, 1926 Venezuela; Colombia; Mexico Eulimnadia texana Packard, 1871 Mexico; Brazil (and North America) Eulimnadia sp. (in Margalef, 1961) Venezuela * Several additional unnamed species exist in private collections; see text its carapace than that of E. belki, Eulim­ If the above differences were the only dis­ nadia brasiliensis differs from E. belki in tinguishing characters I would indeed be that the rostrum of the female is also pro­ hesitant to establish a new species, as so duced, and the frontal organ does not ap­ many characters of the family are variable pear to arise from as marked an elevation (e.g., see Straskraba, 1965). Fortunately, an as seen in E. belki. Unfortunately, speci­ additional character argues strongly for dis­ mens of E. brasiliensis at the Hungarian tinctness of E. belki. The stout, cylindrical Museum of Natural History (see Materials eggs bearing longitudinal grooves and hav­ and Methods) are in very poor condition. ing "flared" rims at either end are appar­ Only the valves and a single egg of one spec­ ently unique in the family. Whereas many imen remain in the vial. Therefore, the other traditionally employed characters of above comparison is based upon the orig­ the Limnadiidae are variable, egg mor­ inal description by Sars (1902). Eulimnadia phology seems to be a conservative char­ chacoensis, which Gumey (1931) felt dif­ acter. As an example, populations of Eulim­ fered "scarcely at all" from E. brasiliensis, nadia in North America that vary in other has an acute male rostrum (although not as characters nevertheless share nearly iden­ pronounced as in E. belki) and very similar tical egg morphology (see Belk, 1989). De­ male claspers. Finally, E, geayi differs from spite the fact that egg morphology was oc­ E. belki not only in size (the carapace of the casionally used to define species of largest specimen of the Daday collections Eulimnadia by Daday as early as 1926, egg in the Hungarian Museum is 10.7 mm long morphology has only infrequently and rel­ and 7.8 mm tall; compared to 3.75 x 2.38 atively recently been employed in compar­ mm for the holotype of E. belki) but also ative studies of branchiopods (e.g., Alonso in having the female rostrum produced. A andAlcaraz, 1984; Belk, 1989; Mura, 1986; vial containing specimens labeled E. co­ Mura and Thiery, 1986; Samyiah et al, lumbica exists in the Collectio Dadayana 1985; Thiery and Champeau, 1988). Phyllopoda of the Hungarian Museum of Of the species of Eulimnadia known from Natural History (see Materials and Meth­ Central and South America (Table 1), there ods); these specimens are also large, ap­ are no species with eggs similar to those of proximately the same size as E. geayi (the E. belki. Eulimnadia brasiliensis, as illus­ largest "£". columbica" specimen was 10.5 trated by Sars (1902), has spherical eggs X 7.1 mm), and the two species are herein bearing smooth bumps on the surface. The considered synonymous (see below). vial containing the single specimen of that

Fig. 5. Various specific characters of species in the Limnadiidae. A, caudal region oi Eulimnadia antillarum (compare reduced posterior spines (small arrows) to Fig. 3F, which is typical of the genus), x 150; B, cylindrical egg (damaged) of £. chacoensis, x350; C, sharp crenulated ridges of egg of is. chacoensis (see arrow in B), X 1,000; D, spherical eggs oi Eulimnadia sp. A [DB 305] from Argentina, x 180; E, Eulimnadia sp. B [DB 632], from Paraguay, showing position of egg mass on mature female; F, higher magnification of eggs shown in E, x75; G, same, x 2,000; H, eggs oi Limnadia lenticularis frompopulatio n south of Tallahassee, Florida, x 270. 112 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 1, 1989 species in the Hungarian Museum of Nat­ egg selected for SEM study cracked badly ural History (see Materials and Methods) upon drying, but the ridges can be seen to contained a single egg that, when subjected be fairly sharp (compared to those of E. to SEM, agrees fairly closely with Sars' il­ belki) and slightly crenulate, a condition that lustration; the egg is roughly spherical with is not brought on by drying conditions (Fig. smooth ridges surrounding irregular depres­ 5B, C). Daday (1926, fig. 138e) described sions (Fig. 4C). Margalef (1961) illustrated the eggs ofE. texana, a predominantly North spherical eggs for "Eulimnadia sp.," but it American species that has been reported is possible that the illustrations were of the from Mexico, as having spherical eggs with "ends" of the eggs and do not show the small protuberances that give the eggs a stel­ cylindrical nature that some other limna- late appearance. Daday's figures are inac­ diid eggs possess; that species must remain curate, since eggs of £". texana are stout and unnamed for the present. Daday's (1926) cylindrical (see Belk, 1989), similar to what illustration of the egg ofE. geayi also shows is described below for an undescribed Par­ a spherical egg with somewhat acute surface aguayan species (DB 632). projections, but this is inconsistent with eggs An examination of the eggs of several un­ o(E. geayi in the Hungarian Museum. The described species, of which larger samples eggs ofE. geayi are short, grooved cylinders exist, serves to illustrate the range of egg with one end of the cylinder slightly wider morphology in the genus. Of the unde­ than the other. The wider end of the cylinder scribed species in the private collections of bears raised ridges, whereas these ridges on Denton Belk, specimens of Eulimnadia sp. the narrow end are flattened or slightly con­ A [DB 305], from Argentina, have spherical cave, giving the egg a somewhat pentagonal eggs with small oval depressions (Fig. 5D). appearance when viewed from the side (Figs. Finally, specimens of the Paraguayan Eu­ 4D-F). The eggs oiE. "columbica"^ (see Ma­ limnadia sp. B [DB 632] have roughly cy­ terials and Methods) are essentially identi­ lindrical eggs that lack flared rims and that cal to those of £•. geayi, confirming that this have deep grooves separated by low, round­ form is a synonym of £•. geayi. Baird (1852, ed longitudinal ridges (Fig. 5E-G); these eggs paper repeated verbatim in 1854) did not are very similar to those known from the illustrate the eggs ofE. antillarum, and un­ predominantly North American E. texana fortunately all of the specimens in the Brit­ (see Belk, 1989). ish Museum (Natural History), and also These and other observations (see Belk, those in the Hungarian Museum, lack eggs. 1989) suggest that egg morphology is species Lilljeborg (1889: 424) stated only "Eggs specific in the genus Eulimnadia and pos­ nodulose or angulose, with 8-10 evident sibly throughout the family Limnadiidae. nodes or angles" (English translation). For­ For example, the distinctive "twisted" eggs tunately, the posterodorsal border of the of North American populations of Limna- caudal region in E. antillarum is distinctive dia lenticular is (Fig. 5H) are identical to in lacking the acute and well-developed those of European populations (compare Fig. spines typical of most other species (the 5H to Sars, 1896). A detailed description of spines are present in some specimens but egg morphology, preferably by means of are very small and low; Fig. 5 A, compare SEM, should be included in any description to Fig. 3F); thus, identification is not diffi­ or redescription of species of the Limna­ cult. However, Daday's illustration (his fig. diidae. In addition, size of the eggs may 127d) of E. antillarum shows more devel­ prove to be of systematic value, although oped spines than are seen in Fig. 5A, and that character was not specifically addressed several authors have noted the variability in this paper. of these spines in the family. Therefore, per­ haps the validity of Eulimnadia antillarum Many authors (see discussion in Webb should be questioned until egg morphology and Bell, 1979) have commented on the is known. Gumey (1931) did not illustrate rather unsatisfactory characters employed the eggs of E. chacoensis, but the eggs of in distinguishing between Eulimnadia and syntypes at the British Museum appear short Limnadia, and in the most recent work ad­ and cylindrical with one end bearing raised dressing the two genera (Webb and Bell, ridges, much as is seen in E. geayi. The one 1979) they were synonymized, Limnadia Brongniart being the senior synonym. I have MARTIN: NEW EVLIMNADIA FROM COEUMEL, MEXICO 113 elected to recognize the two genera on the Fryer, G. 1987. A new classification of the branchio- basis of several morphological characters of pod Crustacea.—Zoological Journal of the Linnean Society 91: 357-383. the telson not illustrated by previous work­ Gumey, R. 1931. Reports of an expedition to Brazil ers (see Belk, 1989). These characters will and Paraguay in 1926-7, supported by the Trustees be addressed in a separate paper. of the Percy Sladen Memorial Fund and the Exec­ utive Committee of the Carnegie Trust for Scotland. ACKNOWLEESGEMENTS Branchiopoda.—Journal of the Linnean Society of London, Zoology 37: 265-275. I am very grateful to Jill Yager for bringing these Hartland-Rowe, R. 1982. , , and specimens to my attention and for providing infor­ Conchostraca.—/n; S. H. Hurlbert, and A. Villalo- mation on the habitat, and to Dr. Denton Belk for bos-Figueroa, eds.. Aquatic biota of Mexico, Central confirming that the species was undescribed. The im­ America and the West Indies. Pp. 175-176. San Di­ portance of egg morphology in conchostracan taxon­ ego State University Foundation, San Diego. omy was brought to my attention by Dr. Belk. I also Lilljeborg, W. 1889. Diagnosen zweier Phyilopoden- thank Dr. Laszlo Forro for help with the literature and Arten aus Siid-Brasilien.-Abhandlungen heraus- for loaning specimens ofE. geayi and the undescribed gegeben vom naturwissenschaftlichen Verein zu Bre­ Colombian Eulimnadia from the Hungarian Museum men 10: 424-457. of Natural History, Dr. Geoffrey Boxshall for loaning Loffler, H. 1977. Notostraca and Conchostraca. — In: E, chacoensis and E. antillarum from the British Mu­ H. S. Hurlbert, ed., Aquatic biota of southern South seum (Natural History), and Mr. Guido Pereira for America. Pp. 131-132. San Diego State University helpful discussion concerning Venezuelan populations. Foundation, San Diego. I am very grateful to Dr. Denton Belk for critical read­ . 1981. Notostraca and Conchostraca. — In: S. ing of the manuscript Special thanks go to Kim Riddle H. Hurlbert, G. Rodriguez, and N. D. Dos Santos, and Sandra Silvers of the Florida State University Elec­ eds., Aquatic biota of tropical South America, part tron Microscopy Center for expert assistance with SEM. 1. Arthropoda. Pp. 3-4. San Diego State University Dr. Lawrence G. Abele provided laboratory space and Foundation, San Diego. facilities during early phases of the study. This work was funded by the National Science Foundation, grant Lutz, A. 1929. Dous phyllopodos, observados no Rio no. BSR-8615018, to J. W. Martin and L. G. Abele. Grande do Norte. —Instituto Oswaldo Cruz, Sup- plemento das Memorias 5: 3-9. Margalef R. 1961. La vida en los charcos de agua LITERATURE CITED dulce de Nueva Esparta (Venezuela).—Memoria de la Sociedad de Ciencias Naturales La Salle 21: 75- Alonso, M., and M. Alcaraz. 1984. Huevos resis- 110. tentes de crustaceos eufilopodos DO cladoceros de la Moore, W. G. 1965. New records of Conchostraca peninsula Iberica: observacion de la morfologia ex­ (Crustacea, Branchiopoda), with an account of their terna mediante tecnicas de microscopia eiectronica occurrence in Louisiana.—Proceedings of the Lou­ de barrido.—Oecologia Aquatica 7: 73-78. isiana Academy of Sciences 28: 41-44. Baird, W. 1852. Monograph of the family Branchi- Mura, G. 1986. SEM morphological survey on the podidae, a family of belonging to the egg shell in the Italian anostracans (Crustacea, Bran­ division Entomostraca, with a description of a new chiopoda).—Hydrobiologia 134: 273-286. genus and species of the family, and two new species , and A. Thiery. 1986. Taxonomical signifi­ belonging to the family Limnadiadae.—Proceedings cance of scanning electron microscopic morphology of the Zoological Society of London 20; 18-30. (Re­ of the euphyllopods' resting eggs from Morocco. Part peated verbatim in 1854, in Annals and Magazine I. Anostraca. —Vie et Milieu 36: 125-131. of Natural History, Second series, 14: 216-229.) Pennak, R. 1978. Freshwater invertebrates of the Belk, D. 1982. Branchiopoda.-Zn.'S. P. Parker, ed.. United States, 2nd edition.—John Wiley and Sons, Synopsis and classification of hving organisms. 2: New York. Pp. i-xv, 1-803. 174-180. McGraw-Hill Book Company, New York. Samyiah, N., K. Venkataraman, and S. Krishnaswamy. . 1989. Identification of species in the con­ 1985. Morphology of three species of Conchostraca chostracan genus Eulimnadia by egg shell morphol­ using scanning electron microscope.—Current Sci­ ogy.—Journal of Crustacean Biology 9: 115-125. ence 54: 869-871. Bowman, T. E. 1971. The case of the nonubiquitons Sars, G. O. 1896. Phyllocarida and Phyllopoda.— telson and the fraudulent furca.—Crustaceana 21: Fauna Norvegiae, 1: 1-140. Joint-Stock Printing 165-175. Company, Christiania. Daday, E. 1926. Monographic systematique des . 1902. 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