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2018 Northeastern Naturalist Notes Vol. 25, No. 2
R.E. Schmidt, E. Kiviat, N. Trigoboff, and J. Vanek
New Records of Clam Shrimp (Laevicaudata, Spinicaudata) from
New York
Robert E. Schmidt1,*, Erik Kiviat1, Norm Trigoboff 2, and John Vanek3, 4
Abstract - We present records of 3 clam shrimp species from New York: 2 spinicaudatan species—
Eulimnadia agassizii (Agassiz Clam Shrimp; the first record for the state) and Cyzicus sp. (the second
record of the genus from the state)—and Lynceus brachyurus (Laevicaudata; Holarctic Clam Shrimp;
first reported from New York in 1883, is herein documented from the Hudson Valley). Some of the
temporary waters that these species inhabit are protected by wetland regulations, but rain-puddle
habitat on or along unimproved roads has no such protection.
Introduction. Clam shrimp are branchiopod crustaceans that generally live in temporary
waters (Williams 1987). The current classification places clam shrimp in 2 orders: the
Laevicaudata and the Diplostraca (suborders Spinicaudata and Cyclestherida, respectively)
(Martin and Davis 2001, Rogers 2009). Two species of clam shrimp are reported in published
literature from New York State—Cyzicus gynecia (Mattox) (Mattox Clam Shrimp)
(Schmidt and Kiviat 2007) and an old record of Lynceus brachyurus Müller (Holarctic
Clam Shrimp) from Long Island (Packard 1883). This note documents 2 more spinicaudatan
species from New York and confirms the presence of L. brachyurus from recently
collected specimens.
Methods. We collected clam shrimp by hand and with a variety of small-mesh dip nets.
We did not collect the specimens as part of targeted searches; rather, we encountered them
during other research activities. We preserved the specimens in 70% isopropanol or 70%
ethanol, identified them under a dissecting microscope with incident and transmitted light,
and made measurements with dividers or an ocular micrometer in a dissecting microscope.
We followed the taxonomic treatments of Smith (1995) and Rogers and Hann (2016).
Results. We report collecting specimens of the following species.
Laevicaudata
Lynceus brachyurus Müller (Holarctic Clam Shrimp). This species is Holarctic in distribution
(Martin and Belk 1988, Rogers and Olesen 2016). In North America, specimens
have been collected from Alberta, Ontario, Quebec, Yukon Territory, Alaska, Arizona, California,
Illinois, Indiana, Massachusetts, Michigan, Minnesota, New Hampshire, New York,
Ohio, Oregon, Rhode Island, and Washington (Martin and Belk 1988), British Columbia
(E-Fauna BC 2016), Montana, and Saskatchewan (Rogers et al. 201 5a).
Packard (1883) reported this species from Glendale, Long Island, NY (as Limnetis
gouldii Baird, a junior synonym; Martin and Belk 1988). There are no published reports of
Holarctic Clam Shrimp from New York in the 130 y since Packard’s (1883) report.
J.H. Bonitatibus observed specimens from temporary pools on 7 June 2014 from 2
locations, both in New York: Dutchess County: Town of Rhinebeck: (1) Stone Church
Road, vernal pool near landfill, 41°57'56.1"N, 73°51'55.9"W; J.H. Bonitatibus; 5 females,
1Hudsonia Ltd., PO Box 5000, Annandale, NY 12504. 2Plant Pathology Herbarium, 334 Tower Road,
Cornell University, Ithaca, NY 14853. 3Department of Biology, Hofstra University, Hempstead, NY
11549. 4Current address - Department of Biological Sciences, Northern Illinois University, DeKalb,
IL 60115. *Corresponding author - schmidt@simons-rock.edu.
Manuscript Editor: David Frings
Notes of the Northeastern Naturalist, Issue 25/2, 2018
2018 Northeastern Naturalist Notes Vol. 25, No. 2
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R.E. Schmidt, E. Kiviat, N. Trigoboff, and J. Vanek
9 males. Specimens ranged from 3.2 mm to 4.0 mm in length. Specimens were deposited
in the National Museum of Natural History (USNM 1454926); and (2) Hook Road, vernal
pool, 41̊°7'46.4"N, 73°54'31"W; J.H. Bonitatibus; no material collected.
Spinicaudata
Cyzicus sp. The genus Cyzicus is in desperate need of revision. Recent preliminary
molecular studies did not support traditionally recognized species, and cryptic species may
have been indicated (Schwentner et al. 2009). This taxonomic uncertainty and revisionary
efforts that are currently underway (Rogers 2016) prevent us from assigning our specimens
to any described species at this time. This genus has a very wide distribution in North
America, from Mexico north to Canada (Smith and Gola 2001), in Alberta, Manitoba,
and Saskatchewan (Williams 1987). In the USA, the range extends from California and
Washington east to Pennsylvania (Wolfe 1982) and Maryland (Smith and Gola 2001), and
southeast to Alabama (Frings and Morse 2015).
The specimens reported here, collected in the vicinity of Ithaca, NY, extend the published
range of this genus about 220 km northward (Wolf 1982) and about 215 km WNW (Schmidt
and Kiviat 2007) of the nearest Cyzicus populations, and are the second record of the genus
from the state: USA: New York: Tompkins County: Town of Danby: Danby State Forest:
rainwater pools on Bald Hill Road, between Station Road and Michigan Hollow Road,
42°17'6"N, 76°29'34"W; 5 September 2016; N. Trigoboff and G. LaBranch; 10 females, 3
males. This site was revisited on 19 July 2017 and the habitat had been destroyed by roadmaintenance
activities. Town of Danby: Danby State Forest: rainwater pools on Bald Hill
Road, between Station Road and Michigan Hollow Road, 42°18'57.3''N, 76°29'41.9''W; 19
July 2017; N. Trigoboff; 14 females, 9 males. Town of Caroline: Shindagin Hollow State Forest:
rainwater pool on jeep trail, 42°20'1"N, 76°20'25"W; 10 September 2016; N. Trigoboff
and G. LaBranch; 14 females. The material ranged in length from 5.6 mm to 12.2 mm. Specimens
were donated to D. Christopher Rogers, University of Kansas, Lawrence, KS.
Eulimnadia agassizii (Packard) (Agassiz Clam Shrimp). This species has been reported
from Massachusetts, Virginia (National Museum of Natural History invertebrate
collection, #62654), the Florida panhandle (Rogers et al. 2010), and Ontario (2 records;
Williams 1987).
We present the first records of this species from New York: Nassau County: Town of
Hempstead: temporary pool on west end of Jones Beach Island, 40°35'22.3"N 73°32'22.7"W;
3 June 2013; J. Vanek; 1 female. Specimen stored at Hudsonia, Ltd., Ulster County: Town of
Ulster: temporary pool on the edge of an organic farm field, 650 m northeast of the junction
of Esopus Avenue and Orlando Street, 41°57'36''N, 74°00'15''W; 17 August 2016; E. Kiviat;
7 ovigerous females. Specimens were deposited in the National Museum of Natural History
(USNM 1454927).
The Jones Beach Island site is in a disturbed area of the maritime dunes ecological community
(Edinger et al. 2014) and is characterized by short, emergent vegetation and an open
canopy. The specimen is 10.1 mm long. The Ulster County pool material ranged in length
from 5.0 mm to 9.1 mm.
Identification of our specimens was based on morphology of the whole organism, which
can be very plastic (Rogers et al. 2012), and egg morphology (Belk 1989, Rogers 2016).
The New York specimens had 4–6 growth lines (Pennak [1989] said ≤4 growth lines, Smith
[1992] indicated 4–5 growth lines). Smith (1992) illustrated a bluntly pointed rostrum, but
the New York material has a round rostrum, and the New York specimens have 12 spines on
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2018 Northeastern Naturalist Notes Vol. 25, No. 2
R.E. Schmidt, E. Kiviat, N. Trigoboff, and J. Vanek
the telson, whereas Smith (1992) gave a range of 11–17. The specimens were not appropriately
preserved for molecular analysis.
Discussion. With the exception of the Rhinebeck specimens, which were found in intermittent
woodland amphibian breeding pools, we found all of the specimens reported here in
small, ephemeral rain pools. Small-puddle inhabitants must have wide tolerance for water
quality, but also for rapid changes in water quality as puddles fill and dry (Rogers 2009).
Some of the extremes that puddle inhabitants survive have been documented (Horne 1971,
Machado et al. 1999, Orridge 2011). Although similar pools on dirt roads and all-terrain–vehicle
trails are common in New York, we have sampled many without finding clam shrimp,
suggesting that the animals are active for very short time-periods, have subtle habitat requirements,
or lack effective dispersal mechanisms.
None of the populations that we report in this note were found as part of a targeted survey,
but rather, we observed them during field work for other purposes. Surveys dedicated
to finding clam shrimp populations would undoubtedly document many more populations
and may well document species not yet known from New York. Although clam shrimp tolerate
various environmental conditions, their habitats, especially rain pools on dirt roads,
are often continually extant for only relatively short periods of time. No laws protect these
habitats. Large branchiopods worldwide are diverse and poorly known biogeographically
and ecologically; many species have been described only recently. There are many narrowly
endemic species, and many taxa are threatened by human activities (Rogers et al.
2015b, Schmidt and Kiviat 2007). We recommend additional survey efforts to determine
the distribution and conservation needs of these species and their habitats in New York
and adjacent areas.
Acknowledgments. We thank Creek Iverson for facilitating field work on the Northeast Farms
site in Ulster County and Jill Bonitatibus Pritchett for collecting Lynceus from Rhinebeck. We also
thank The Ithaca Hikers for their assistance. The manuscript was improved with the help of David
Frings, D. Christopher Rogers, and an anonymous reviewer. This paper is a Bard College–Hudsonia
contribution.
Literature Cited
Belk, D. 1989. Identification of species in the conchostracan genus Eulimnadia by egg-shell morphology.
Journal of Crustacean Biology 9:115–125.
Edinger, G.J., D.J. Evans, S. Gebauer, T.G. Howard, D.M. Hunt, and A.M. Olivero. 2014. Ecological
Communities of New York State, 2nd Edition. A revised and expanded edition of Carol Reschke’s
Ecological Communities of New York State. New York Natural Heritage Program, New York State
Department of Environmental Conservation, Albany, NY. 160 pp.
E-Fauna BC. 2016. Electronic atlas of the wildlife of British Columbia. Available online at http://
linnet.geog.ubc.ca/efauna/Atlas/Atlas.aspx?sciname=Lynceus%20brachyurus. Accessed 3 November
2016.
Frings, D.M., and K.J. Morse. 2015. New state record and southeastern United States range extension
for Cyzicus mexicanus (Claus) (Mexican Clam Shrimp). Southeastern Naturalist 14:N67– N69.
Horne, J.H. 1971. Some effects of temperature and oxygen concentration on phyllopod ecology. Ecology
52:343–347.
Machado, M., M. Cristo, and D.C. Da Fonseca. 1999. Non-cladoceran branchiopod crustaceans from
southwest Portugal. I. Occurrence notes. Crustaceana 72:591–602 .
Martin, J.W., and D. Belk. 1988. Review of the clam shrimp family Lynceidae Stebbing, 1902 (Branchiopoda:
Conchostraca) in the Americas. Journal of Crustacean Biology 8:451–482.
Martin, J.W., and G.E. Davis. 2001. An updated classification of the recent Crustacea. Natural History
Museum of Los Angeles County, Science Series 39:124 pp.
2018 Northeastern Naturalist Notes Vol. 25, No. 2
N10
R.E. Schmidt, E. Kiviat, N. Trigoboff, and J. Vanek
Orridge, J. 2011. Genetic, morphological and ecological relationships among populations of the clam
shrimp, Caenestheriella gynecia. Ph.D. Dissertation. City University of New York, New York,
NY. 185 pp.
Packard, A.S., Jr. 1883. A monograph of the phyllopod Crustacea of North America, with remarks on
the order Phyllocarida. Annals and Magazine of Natural History 12:295–419.
Pennak, R.W. 1989. Fresh-water Invertebrates of the United States. John Wiley and Sons, New York,
NY. 628 pp.
Rogers, D.C. 2009. Branchiopoda (Anostraca, Notostraca, Laevicaudata, Spinicaudata, Cyclestherida).
Encyclopedia of Inland Waters 2:242–249.
Rogers, D.C., and B.J. Hann. 2016. Class Branchiopoda. P p. 437–476. In J.H. Thorp and D.C. Rogers
(Eds.). Thorp and Covich’s Freshwater Invertebrates. Keys to Nearctic Fauna. Academic Press,
Cambridge, MA. 762 pp.
Rogers, D.C., and J. Olesen. 2016. Laevicaudata catalogus (Crustacea: Branchiopoda): An overview
of diversity and terminology. Arthropod Systematics and Phylogeny 74:221–240.
Rogers, D.C., S.C. Weeks, and W.R. Hoeh. 2010. A new species of Eulimnadia (Crustacea: Branchiopoda:
Diplostraca: Spinicaudata) from North America. Zootaxa 2413:61–68.
Rogers, D.C., N. Rabet, and S.C. Weeks. 2012. Revision of the extant genera of Limnadiidae (Branchiopoda:
Spinicaudata). Journal of Crustacean Biology 32:827–8 42.
Rogers, D.C., J. Olesen, and J.W. Martin. 2015a. A new possibly parthenogenic species of Lynceus
from Canada (Crustacea: Branchiopoda: Laevicaudata), with key to the Nearctic female Laevicaudata.
Scientific Papers, Natural History Museum, The University of Kansas 47:1–9.
Rogers, D.C., M. Schwentner, J. Olesen, and S. Richter. 2015b. Evolution, classification, and global
diversity of large Branchiopoda. Journal of Crustacean Biology 35:297–300.
Schmidt, R.E., and E. Kiviat. 2007. State records and habitat of clam shrimp, Caenestheriella gynecia
(Crustacea: Conchostraca), in New York and New Jersey. Canadian Field-Naturalist 121:128–132.
Schwentner, M., B.V. Timms, R. Bastrop, and S. Richter. 2009. Phylogeny of Spinicaudata (Branchiopoda,
Crustacea) based on three molecular markers: An Australian origin for Limnodopsis.
Molecular Phylogenetics and Evolution 53:716–725.
Smith, D.G. 1992. A redescription of types of the Clam Shrimp Eulimnadia agassizii (Spinicaudata:
Limnadiidae). Transactions of the American Microscopical Society 111:223–228.
Smith, D.G. 1995. Keys to the Freshwater Macroinvertebrates of Massachusetts, 2nd Edition. University
of Massachusetts, Biology Department, Amherst, MA. 236 pp.
Smith, D.G., and A.A. Gola. 2001. The discovery of Caenestheriella gynecia Mattox 1950 (Branchiopoda,
Cyzicidae) in New England, with ecological and systematic notes. Northeastern Naturalist
8:443–454,
Williams, D.D. 1987. The Ecology of Temporary Waters. Timber Press, Portland, OR. 205 pp.
Wolfe, A.F. 1982. Distribution of Cyzicus mexicanus (Conchostraca: Crustacea) in Lebanon County,
Pennsylvania. Proceedings of the Pennsylvania Academy of Science 56:36–38.