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Untangling Confusion Between Eubranchipus Vernalis and Eubranchipus Neglectus (Branchiopoda: Anostraca) Author(S): Denton Belk, Graziella Mura, Stephen C

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Untangling Confusion Between Eubranchipus Vernalis and Eubranchipus Neglectus (Branchiopoda: Anostraca) Author(S): Denton Belk, Graziella Mura, Stephen C Untangling Confusion between Eubranchipus vernalis and Eubranchipus neglectus (Branchiopoda: Anostraca) Author(s): Denton Belk, Graziella Mura, Stephen C. Weeks Source: Journal of Crustacean Biology, Vol. 18, No. 1 (Feb., 1998), pp. 147-152 Published by: The Crustacean Society Stable URL: http://www.jstor.org/stable/1549529 Accessed: 11/11/2010 16:16 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. 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The Crustacean Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Crustacean Biology. http://www.jstor.org JOURNALOF CRUSTACEANBIOLOGY, 18(1): 147-152, 1998 UNTANGLING CONFUSION BETWEEN EUBRANCHIPUS VERNALISAND EUBRANCHIPUSNEGLECTUS (BRANCHIOPODA: ANOSTRACA) Denton Belk, Graziella Mura, and Stephen C. Weeks ABSTRACT Despiteillustrations in Garman(1926) clearlyshowing the differentantennal appendages of Eu- branchipus vernalis and Eubranchipus neglectus, Creaser (1930) published erroneous drawings of the antennalappendages of these two species thatled to more than65 years of taxonomicconfu- sion betweenthem. We untanglethis confusion,and show thatthese species have nonoverlapping areasof occurrencewith E. vernalisto the eastand E. neglectusto the westof the AppalachianMoun- tains.In addition,we presentevidence supporting the use of resting-egg(cyst) morphologyin study- ing evolutionaryrelationships among anostracan species. An importantpart of this usefulnessis due to the independenceof cyst morphologyfrom sexual selection.Since the primarytaxonomic char- actersof anostracansare all stronglyinfluenced by sexualselection, cyst morphologycan supplya reasonablyindependent set of charactersfor testinghypotheses of speciesrelationships. Nine of the 16 anostracanspecies described similarity,a situation that usually indicates in the genus Eubranchipusare endemic to close relationship. North America (Belk and Brtek, 1995). Of Brendonck et al. (1992) found that four these nine, two species pairs, bundyi-intrica- Africanspecies in the anostracangenus Strep- tus and holmanii-moorei,were involved in tocephalusproduce uniquely shaped tetrahe- taxonomic problemsthat have alreadybeen dral cysts. Their morphologicalanalysis of cleared up. Here we reveal and correctcon- the taxonomicallyimportant male antennae fusioninvolving a thirdspecies pair, vernalis- suggestedthese four streptocephalidsform a neglectus.In addition,we show thatthe mor- closely relatedgroup. Thus, as it turnedout, phology of the cyst shell reflects the appar- the unique tetrahedralcyst representedthe ent close evolutionaryrelationship between most striking of a cluster of morphological the sister species of each pair. charactersindicating close evolutionaryre- Hartland-Rowe(1967) pointedout thattwo lationship.Brendonck et al. formallyrecog- taxa were being confusedunder the nameEu- nized this situationby assigningthe four spe- branchipus bundyi Forbes, 1876. He docu- cies to a new subgenus, Parastreptocephalus. mented the morphologicaldifferences, and This is the first example of cyst morphology described a new species, Eubranchipus intri- aiding in the discovery of a groupof closely catus Hartland-Rowe, 1967. Brtek (1967) relatedanostracans. demonstratedthat the taxon from southern As was the case in the groupof relatedspe- Louisianareferred to in severalstudies by Dr. cies which Brendoncket al. (1992) named Walter G. Moore as Eubranchipus holmanii Parastreptocephalus, we found each of the (Ryder,1879) was in fact a new specieswhich threepairs of problemspecies of Eubranchi- he named Eubranchipus moorei Brtek, 1967. pus producecysts that look alike in compar- We point out below that a mistake in fig. 3 isons made between the members of each of Creaser(1930) resulted in confusion be- pair.The threespecies that have not been con- tween Eubranchipus vernalis (Verrill, 1869) fused with other taxa (Eubranchipus ore- and Eubranchipus neglectus Garman, 1926. gonus Creaser, 1930, Eubranchipus ornatus Creaser'serror was picked up and perpetu- Holmes, 1910, and Eubranchipus serratus ated in popular identificationkeys by Pen- Forbes, 1876) producecysts that look differ- nak (1953, and later editions) and by Dexter ent from each other,and from cysts of all the (1959). Brtek(1966) incorrectlyplaced these other North Americanspecies. two in synonymy,while hesitantlysuggesting the neglectus form might be a subspecies. MATERIALSAND METHODS Confusion between the taxa within each of All cystsused in thisstudy were removed from the brood these threepairs resulted from morphological pouchesof preservedspecimens. We choseonly cysts that 147 148 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 18, NO. 1, 1998 Figs. 1, 2. Dorsal views of left antennal appendages. 1, Eubranchipus neglectus, antennal appendage from a 25-mm male collected in Cuyahoga County, Ohio, U.S.A. (DB146); 2, Eubranchipus vernalis, antennal appendage from a 20-mm male collected in New London County, Connecticut, U.S.A. (DB1095). Scales = 1 mm. appearedto be mature and of normal morphology, so as to male by SEM. When materialwas available, we used cysts avoid problems like those discussed in Mura (1992). The from several populations, choosing especially those distant selected cysts were preparedfor SEM analysis as described from each other. For our figures, we selected the SEM in Mura (1986). We used cysts of 1-5 females from each photographs that most accurately illustrated the typical location, and examined more than 4 cysts from each fe- morphology of the cyst produced by each species. Figs. 3, 4. Cysts, whole view (a) and close view of surface (b). 3, Eubranchipus holmanii; 4, Eubranchipus moorei. Scales: a = 100 am; b = 20 ,um. BELK ETAL.: SYSTEMATICS AND CYST MORPHOLOGY OF EUBRANCHIPUS 149 Figs. 5-8. Cysts, whole view (a) and close view of surface (b). 5, Eubranchipus bundyi; 6, Eubranchipus intricatus; 7, Eubranchipus vernalis; and 8, Eubranchipus neglectus. Scales: a = 200 lim; b = 50 jim. 150 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 18, NO. 1, 1998 Figs. 9-11. Cyst, whole view (a) and close view of surface (b). 9, Eubranchipus ornatus; 10, Eubranchipus serra- tus; and 11, Eubranchipus oregonus. Scales: a = 100 gm; 9b = 20 gm; 10b and lb = 50 gm. We used cysts from the following sources and locali- serratus: DB152 Coconino County, Arizona, U.S.A.; ties during the SEM portion of our study. For E. bundyi: DB581 British Columbia, Canada; and DB663 Beaver- DB222 (DB = collection of Denton Belk) Apache County, head County, Montana, U.S.A. For E. vernalis: DB 1095 Arizona, U.S.A.; DB229 Coconino County, Arizona, New London County, Connecticut, U.S.A. and YPM8247 U.S.A.; DB528 Alberta, Canada; and DB792 Northwest (YPM = Peabody Museum of Natural History, Yale Uni- Territories, Canada. For E. holmanii: DB517 Madison versity) New Haven County, Connecticut, U.S.A. County, Alabama, U.S.A. For E. intricatus: DB554 Al- Data for the map in Fig. 12 came from field work by berta, Canada, and DB557 Alberta, Canada. For E. Stephen Weeks in Ohio during the period 26 July 1995 moorei: DB521 Mobile County, Alabama, U.S.A., and and 1 May 1996, and locality data on collections in the DB524 (paratypes) St. Tammany Parish, Louisiana, holdings of the National Museum of Natural History, U.S.A. For E. neglectus: DB 146 Cuyahoga County, Ohio, Smithsonian Institution; the Milwaukee Public Museum, U.S.A.; DB518 Madison County, Alabama, U.S.A.; Milwaukee, Wisconsin, U.S.A.; the Peabody Museum of DB880 Madison County, Alabama, U.S.A.; and accession Natural History, Yale University; the Canadian Museum no. IZ1994-12 (Milwaukee Public Museum, Milwaukee, of Nature, Ottawa, Ontario; the personal collection of Wisconsin, U.S.A.) Jefferson County, Kentucky, U.S.A. Denton Belk; and specimens examined by Belk for Harp For E. oregonus: DB727 King County, Washington, et al. (in press). These specimens are divided among the U.S.A. For E. ornatus: DB530 Alberta, Canada. For E. Smithsonian Institution, Arkansas State University Mu- BELK ETAL.: SYSTEMATICS AND CYST MORPHOLOGY OF EUBRANCHIPUS 151 vernalis by comparing Figs. 7 and 8. In ad- dition, very similar ridge development, dif- fering only in width, suggested that bundyi, intricatus, neglectus, and vernalis may be a cluster of related species (Figs. 5-8). Each of the three species of Eubranchipus that have not been subjects of taxonomic problems had a unique cyst-shell patternin comparison to all the other North American species (Figs. 9-11). For these three, cyst morphology offered no suggestion of their relationships among the North American
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