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Geographical color pattern of Argia apicalis (Odonata: Coenagrionidae) in the absence of molecular variation.

With only a few exceptions, odonates do not exhibit much intraspecific color pattern variation that is clearly correlated with geography despite distributions that encompass several ecological niches and wide areas. A noted exception is the Argia fumipennis (Odonata: Coenagrionidae) complex found in the southeastern region of the United States, which is composed of 3 sub-species: A. fumipennis fumipennis (Burmeister), A. fumipennis violacea (Hagen), and A. fumipennis atra Gloyd, and the subspecies are distinguished based only on wing color and distribution (Burmeister 1839; Hagen 1861; Gloyd 1968). Argia apicalis (Say) (Coenagrionidae: Odonata) is a species complex that shows a similar geographical color pattern variation.

Argia apicalis is an ecologically vagile species inhabiting both pond and stream environments. Say (1839) diagnosed A. apicalis as having a pearlaceous blue thoracic region and a hairline humeral and thin dorsal stripe (Fig. 1A). Other characters used to identify A. apicalis are the distinctive pointed and tooth-like cercus in males (Fig. 1B) and a distribution east of the Rocky Mountains and into Arizona (Fig. 1C) (Garrison 1994).

Bick & Bick (1965), Dunkle (1990), Johnson & Westfall (1970), and Johnson (1972) noted variation of the humeral stripe in southeastern populations of A. apicalis. The subpopulation of A. apicalis in the southeastern range (Suwannee County and Columbia County, Florida) have a broad humeral stripe across the length of the pterothorax. In his analysis of the geographical variation within A. apicalis in Florida, Johnson (1972) categorized both males and females into groupings based on the amount of black markings on the head, thorax, and abdomen, and defined "typical" A. apicalis as having humeral stripe values of 1 or 2, whereas "atypical" specimens had values of 4 or 5. Specimens with a value of 3 were deemed intermediates. Johnson (1972) then discussed the pattern variability in the southeastern distribution and noted that in this region specimens with the typical pattern were less than 5% and specimens with atypical thoracic patterns represented more than 95% of the population. In the summer of 2013, 2 authors herein (Smith-Herron & T. Cook) collected A. apicalis near the Suwannee River and noted that 100% of their specimens (122 specimens) fit Johnson's definition of atypical. This study is the first to combine color pattern and cytochrome-b gene sequences to document variation within an extensive portion of the distribution range of A. apicalis.

Materials and Methods

INSECT COLLECTIONS

Argia apicalis adults were collected using aerial nets in May through Sep 2013 and 2014 from 10 localities in Texas, Louisiana, Florida, and Oklahoma (Table 1). Individuals were field preserved in 70 to 90% ethanol and subsequently dried and curated upon arrival at the laboratory. Specimens were photographed with a Canon EOS 70D mounted on an Olympus SZX12 microscope.

We also obtained about 200 specimens on loan from the following museum collections: National Museum of Natural History (NMNH), International Odonata Research Institute (IORI), Georgia Museum of Natural History (GMNH), and Sam Houston State Entomology Collection (SHSUEC), which allowed us to augment our coverage of the distributional range of A. apicalis. The combination of our field-collected samples with existing museum specimens represent about 59% of the reported distribution of A. apicalis in the United States (Fig. 2) (Westfall & May 2006).

MORPHOLOGICAL CHARACTERS

Since its description in 1839, researchers have used hairline humeral stripe found on the thorax to identify and distinguish A. apicalis from its congeners (Johnson 1972). We examined the width of the humeral stripe of the thorax and the pattern and coloration of the head and thorax. For the humeral stripe, the width of the stripe, along the entire length of the thorax, was evaluated and whether or how much it narrowed along its length (Fig. 3). Johnson (1972) reported that individuals with large black patterning on the head usually had hairline humeral stripes whereas individuals with small patterning had wider humeral stripes. To test this observation, we examined head and abdomen of specimens for distinct patterns between the southeastern and the northwestern populations. We also evaluated variation in the morphology of the male caudal appendages and female mesostigmal plates (Garrison 1994; Westfall & May 2006).

MOLECULAR ANALYSES

We extracted genomic DNA from legs for a subset of 29 individuals (Table 2) following the standard protocol instructions of Zymo Research's Quick-gDNA[TM] Mini Prep extraction kit (Zymo Research, Irvine, California). Individuals selected for DNA extraction were taken from field-collected specimens as most museum specimens included in the morphological analyses were collected over 20 yr ago. Special care was taken to select individuals from populations throughout the geographic distribution of A. apicalis (i.e., Florida, Kansas, Louisiana, Oklahoma, and Texas). Although the mitochondrial cytochrome oxidase I (COI) gene has been used widely to detect cryptic diversity and inter-population differentiation in odonates (Brown et al. 2000) and other hexapods (Folmer et al. 1994; Marcus et al. 2009), repeated attempts to amplify this gene fragment in our samples using various previously published primers [e.g. HCO/LCO (Folmer et al. 1994), EVA/ JERRY (Blum et al. 2003)] failed to produce positive amplicons. We thus decided to PCR amplify a 361 bp fragment of the mitochondrial gene cytochrome-b (cyt-b) using primers (151F: 5'-TGTGGRGCNACYGTW-3'; 270R: 5'-AANAGGAARTAYCAYTCNGGYTG-3') and conditions published by Merritt et al. (1998). The usefulness of this gene fragment in detecting cryptic diversity and population structure in insects and other arthropods is well established (Simmons & Weller 2001; Santamaria et al. 2013). Positive amplicons were cleaned and sequenced at Genewiz (South Plainfield, New Jersey), with resulting sequences assembled and edited (e.g., removal of primer regions) using Geneious R 8.0.2 (Biomatters Ltd.). Dried specimens from which DNA was extracted were labeled as DNA vouchers and deposited into the SHSUEC. DNA template vouchers are stored at -40 [degrees]C in the Sam Houston State Natural History Collections (SHSNHC) housed at the Texas Research Institute for Environmental Studies (TRIES) facility, Sam Houston State University.

Results

ANALYSIS OF COLOR PATTERN

Specimens from the northwestern distribution of A. apicalis (north of the Florida Panhandle) displayed a "typical" hair-line humeral stripe. The humeral stripe in this distribution range varied from almost absent in some specimens to very narrow at most, and rarely extended more than a third of the length of the pterothorax. Although the thickness of the humeral stripe varied in the northwestern populations, the variation was not as pronounced as that between northwestern and southeastern populations (Figs. 4-6). Southeastern populations had an "atypical" humeral stripe that was much wider than the "typical" variation and generally extended beyond the anterior third of the pterothorax and might extend the entire length of the pterothorax. Patterns on the head and abdomen were variable among all individuals and were not correlated with geography. After examining all specimens, we noted that the black patterns on the head and abdomen were variable across all specimens and all geographic regions.

MOLECULAR ANALYSIS

All sequences produced in this study were deposited in GenBank under accession numbers KP770147 to KP770165. We successfully sequenced the target cyt-b gene fragment for 19 A. apicalis individuals from throughout its range in the United States: 8 from 5 populations in Texas, 7 from a single population in Florida, 3 from 2 populations in Oklahoma, and 1 from a single population in Kansas. All 19 individuals harbored a single haplotype for the cyt-b gene, indicating no divergence between individuals, populations, or regions. The remaining 11 DNA extractions (4 from Texas, 1 from Florida, 1 from Oklahoma, 2 from Louisiana, 2 from Kansas, and 1 from Nebraska) failed to produce positive amplicons despite repeated attempts at PCR amplification and re-extraction of DNA. Given our findings of non-existent genetic divergence in this gene, we decided against further efforts to produce sequences for these samples.

Discussion

The 2 color forms of A. apicalis were described briefly by Johnson (1972) and Dunkle (1990). Descriptions of A. apicalis have characterized it as having a pterothorax with a narrow dorsal stripe and either no or at most thin humeral stripe, except for populations from Florida in which the humeral stripe is wider and extends more than half the length of the pterothorax. The geographical divide of the 2 color forms is the Suwannee River and reflects the population disjunctions of the sub-species of A. fumipennis in which A. f. atra occurs east of the Suwannee River, A. f. fumipennis west of the Suwannee River, and A. f. violacea north/northwest of Florida. Where the color morphs overlap, they are intermediate in the color pattern and cannot be identified further.

The 2 color forms of A. apicalis were once documented to coexist near the Suwannee River but their co-existence may no longer be true. As we found no differences in cyt-b gene sequences between the 2 color morphs throughout their distribution range, their recognition as separate species is not justified. Although no mechanism or cause is known for the variation seen in both the A. fumipennis and A. apicalis complexes, Johnson (1972) suggested allopatric processes associated with sea level changes during the Pleistocene (e.g., fragmentation of Florida into islands and formation of the Suwannee Straits) may be responsible for the observed differences.

Acknowledgments

The authors would like to thank Karen Pruitt and Cowboy Joe Herron for aiding in the collection of specimens from west and south Texas and Lenora Reid for granting permission to collect on her property. We would also like to extend our appreciation to Daniel Haarmann, Sibyl R. Bucheli, Brent C. Rahlwes, Soheyla Bayat, and Ashley R. Morgan for their assistance in the identification of specimens and discussion of molecular techniques. We are truly grateful to Christopher P. Randle, who allowed the use of his research lab to conduct the molecular portion of this research. The authors would like to acknowledge the Georgia Museum of Natural History, the International Odonata Research Institute, the National Museum of Natural History, and the Sam Houston State University Natural History Collections for providing loan specimens that aided in the overall research. Collection permits were granted by the Florida Department of Environmental Protection Division of Recreation and Parks (#05291310) and the Louisiana Department of Wildlife and Fisheries. This research was made possible by funding provided by a Sam Houston State University Enhancement Research Grant (#290041). Lastly, we would like to thank the several anonymous reviewers for their careful edits and helpful comments that greatly improved the manuscript.

References Cited

Bick GH, Bick JC. 1965. Color variation and significance of color in reproduction in the damselfly, Argia apicalis (Say) (Zygoptera: Coenagrionidae). Canadian Entomologist 97: 32-41.

Blum MJ, Bermingham E, Dasmahapatra K. 2003. A molecular phylogeny of the Neotropical butterfly genus Anartia (Lepidoptera: Nymphalidae). Molecular Phylogenetics and Evolution 26: 46-55.

Brown JM, McPeek MA, May ML. 2000. A phylogenetic perspective on habitat shifts and diversity in the North American Enallagma damselflies. Systematic Biology 49: 697-712.

Burmeister H. 1839. Handbuch der Entomologie. Zweiter Band, pp. 757-1050. Theod. Chr. Friedr. Enslin, Berlin, Germany.

Dunkle S. 1990. Damselflies of Florida, Bermuda and the Bahamas. Scientific Publishers, Gainesville, Florida.

Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294-299.

Garrison RW. 1994. A synopsis of the genus Argia of the United States with keys and descriptions of new species, Argia sabino, A. leonorae, and A. pima (Odonata: Coenagrionidae). Transactions of the American Entomological Society 120: 287-368.

Gloyd LK. 1968. The union of Argia fumipennis (Burmeister, 1839) with Argia violacea (Hagen, 1861), and the recognition of three subspecies (Odonata). Occasional papers of the Museum of Zoology, University of Michigan 658: 1-6.

Hagen H. 1861. Synopsis of the Neuroptera of North America. Smithsonian Institution, Washington, District of Columbia.

Johnson C. 1972. An analysis of geographical variation in the damselfly, Argia apicalis (Zygoptera: Coenagrionidae). The Canadian Entomologist 104: 1515-1527.

Johnson C, Westfall MJ. 1970. Diagnostic keys and notes on the damselflies (Zygoptera) of Florida. Bulletin of the Florida State Museum, Biological Sciences 15: 45-89.

Marcus JM, Bell DD, Bryant AN, Burden EC, Carter ME, Cataldo TJ, Clark KR, Compton HE, DeJarnette LS, Faulkner VB, Gregory RW, Hall JR, Houchin LN, Hudson ME, Jenkins III PF, Jordan JM, Logan BK, Long NR, Maupin HF, McIntyre SR, Mitchell JK, Mobley JK, Nehus AN, Potts BN, Read CR, Slinker KN, Thompson CE, Hughes TM, McElroy DM, Wyatt RE. 2009. The Upper Green River Barcode of Life Project. Journal of the Kentucky Academy of Sciences 70: 75-83.

Merritt TJS, Shi L, Chase MC, Rex MA, Etter RJ, Quattro JM. 1998. Universal cytochrome b primers facilitate intraspecific studies in molluscan taxa. Molecular Marine Biology and Biotechnology 7: 7-11.

Santamaria CA, Mateos M, Taiti S, DeWitt TJ, Hurtado LA. 2013. A complex evolutionary history in a remote archipelago: phylogeography and morphometrics of the Hawaiian endemic Ligia isopods. PLoS One 8: e85199.

Say T. 1839. Descriptions of the new North American neuropterous insects and observations on some already described by (the late) Th. Say. Journal of the Academy of Natural Science of Philadelphia 8: 9-46.

Simmons RB, Weller SJ. 2001. Utility and evolution of cytochrome b in insects. Molecular Phylogenetics and Evolution 20: 196-210.

Westfall MJ, May ML. 2006. Damselflies of North America. Scientific Publishers, Gainsville, Florida.

Melissa S. Sisson (1,2) *, Carlos A. Santamaria (3,4), Autumn J. Smith-Herron (1), Tamara J. Cook (3), and Jerry L. Cook (5)

(1) Texas Invasive Species Institute, Sam Houston State University, Huntsville, Texas 77341, USA; E-mail: melissa.sisson@und.edu (M. S. S.), smith-herron@shsu.edu (A. J. S.-H.)

(2) University of North Dakota, Department of Biology, Grand Forks, North Dakota 58203, USA

(3) Sam Houston State University, Department of Biological Sciences, Huntsville, Texas 77341, USA; E-mail: csantamaria@sar.usf.edu (C. A. S.), bio_tjc@shsu.edu (T. J. C.)

(4) Biology Faculty, College of Arts and Sciences, University of South Florida Sarasota-Manatee, Sarasota, Florida 34243, USA

(5) Sam Houston State University, Office of Research and Sponsored Programs, Huntsville, Texas 77341, USA; E-mail: bio_jlc@shsu.edu (J. L. C.)

* Corresponding author; Email: melissa.sisson@und.edu (M. S. S.)

Caption: Fig. 1. The 4 defining characters of Argia apicalis; (A) a pearlaceous blue pterothorax and a hairline humeral stripe; (B) a thin dorsal stripe; (C) males have a distinctive pointed and tooth-like cercus; (D) and their distribution east of the Rocky Mountains (shaded areas on map are the recorded distribution of A. apicalis).

Caption: Fig. 2. (A) Distribution map of specimens examined (this accounts for about 59% of the reported distribution of Argia apicalis). Dark gray states with black dots represent collected specimen localities, and light gray states with no dots represent areas where no specimens were collected. (B) Distribution of color morphs of A. apicalis in Florida; dark gray represents counties with typical A. apicalis, light gray represents counties with atypical A. apicalis, and the striped area represents the county in which both typical and atypical A. apicalis morphs are present.

Caption: Fig. 3. Geographical color variation: individuals from the (A) northwestern range of the distribution have a typical (= narrow) humeral stripe, whereas individuals from the (B) southeastern part of the range have an atypical (= wide) humeral stripe.

Caption: Fig. 4. Variation in width and extension of humeral stripes in the northwest region of the distribution: (A) Dallas County, Texas; (B) Fairfax County, Virginia; (C) Hunterdon County, New Jersey; (D) Iowa; (E) Missouri County, Oregon; (F) Washington Parish, Louisiana; (G) Holmes County, Florida; (H) Wharton County, Texas; and (I) Washington D.C.

Caption: Fig. 5. Identifying characters of Argia apicalis in the southeast: (A) humeral stripe wide extending at least three-quarters of the pterothorax length; (B) middorsal line slightly wider than in northwestern A. apicalis; and (C) paler caudal appendage (whiter) than in individuals from the north.

Caption: Fig. 6. Variation of humeral stripes in the southeast region of the distribution: (A) Clay County, Florida; (B) Columbia County; Florida; (C) Wakulla County, Florida; and (D) Suwannee County, Florida.

Table 1. List of Argia apicalis specimens examined for
color pattern analysis.

Museum                Date          Sex      State
accession
numbers

USMN 00354813    1 VIII 1914         F        DC
USMN 00354821    22 VIII 1974        M        FL
USMN 00354822    22 VIII 1974        F        FL
IORI 00038262    10 VI 1954         --        FL
IORI             15 VIII 1969        M        FL
IORI             12 IV 1975          F        FL
IORI             25 VIII 1950        M        FL
IORI             12 VII 1972         M        FL
IORI             2 VIII 1969         M        FL
IORI             11 VIII 1969        M        FL
IORI             28 V1973            M        FL
IORI             28 V1973            F        FL
IORI             22 VIII 1984        F        FL
IORI             12 VIII 1969        M        FL
IORI             7 V 1981           --        FL
IORI             33 VIII 1984        M        FL
IORI             12 VII 1972         F        FL
IORI             29 IV 1973          M        FL
IORI             18 IX 1971          F        FL
USMN 00354826    20 VIII 1946       M/F       GA
GMNH             10 VII 1955        --        GA
GMNH             7 VII 1971         --        GA
GMNH             23 VI 1950         --        GA
GMNH             20 V1984           --        GA
GMNH             13 VI 1950         --        GA
GMNH             22-27 VII 1970     --        GA
GMNH             20 VI 1938         --        GA
GMNH             20 VI 1937         --        GA
GMNH             26V1953            --        GA
GMNH             9-13 V 1994        --        GA
GMNH             VI 1937            --        GA
GMNH             2 VII 1944         --        GA
GMNH             27 VII 1944        --        GA
GMNH             2 VII 1944         --        GA
GMNH             25 VII 1941        --        GA
GMNH             24 VII 1931        --        GA
GMNH             16 VI 1946         --        GA
USMN 00354841    24 VII 1929         M        IL
USMN 00354842    24 VII 1929         F        IL
USMN 00355162    26 VII 1953         M        IN
USMN 00355164    26 VII 1953         M        IN
USMN 00354873    8 VII 1927          M        IA
USMN 00354875    15 VII 1927         F        IA
USMN 00354900    10 VII 1943         M        KS
USMN 00354901    10 VII 1943         F        KS
USMN 00354908    18 VI 1947          F        KY
USMN 00354906    29 VII 1948         M        KY
USMN 00354909    12 VIII 1925        F        LA
USMN 00354910    12 VIII 1925        M        LA
USMN 00354930    21-28 VII 1903      M        MD
USMN 00354928    21-28 VII 1903      F        MD
USMN 00354969    16 VII 1907         M        MN
USMN 00354970    16 VII 1907         F        MN
USMN 00354981    28 VIII 1949        F        MO
USMN 00354979    28 VIII 1949        M        MO
GMNH             16 IX 1972         --        MO
USMN 00355192    20 VII 1998         M        NB
USMN 00355192    20 VII 1998         M        NB
USMN 00355200    3 VIII 1985         F        NJ
USMN 00355204    3 VIII 1985         M        NJ
USMN 00355209    19 IX 1988          M        NM
USMN 00355213    18 VIII 1939        F        NC
USMN 00355214    18 VIII 1939        M        NC
USMN 00355218    20 VII 1926         F        OH
USMN 00355222    20 VII 1926         M        OH
USMN 00355247    1962                F        OK
USMN 00355248    1960                M        Ok
USMN 00355254    13-14 VI 1970       M        PA
USMN 00355257    18 VII 1939         M        SC
USMN 00355256    18 VII 1939         F        SC
USMN 00355275    14 VIII 1939        F        TN
USMN 00355272    31 V 1953           M        TN
USMN 00355297    7 IX 1949           M        TX
USMN 00355303    15-17 VI 1965       M        TX
USMN 00355304    8 VI 1965           F        TX
USMN 00355309    26 VI 1904         M/F       TX
USMN 00355306    15-17 VI 1965       M        TX
USMN 00391416    12-26 VII 2007      F        VA
USMN 00391417    12-26 VII 2007      M        VA
USMN 00713680    13 VI 2007          F        WV
USMN 00713676    24 VIII 2004        M        WV
SHSUE 026337     20 VII 2013         M        FL
SHSUE 026336     20 VII 2013         M        FL
SHSUE 026339     20 VII 2013         F        FL
SHSUE 026338     20 VII 2013         M        FL
SHSUE 026335     20 VII 2013         F        FL
SHSUE 026344     20 VII 2013         F        FL
SHSUE 026345     20 VII 2013         M        FL
SHSUE 026349     31 VIII 2013        F        TX
SHSUE 026348     31 VIII 2013        F        TX
SHSUE 026346     31 VIII 2013        M        TX
SHSUE 000185     1 IX 2007          --        OK
SHSUE 000161     23 VII 2007        --        TX
SHSUE 000189     10 VIII 2007       --        TX
SHSUE 001293     25 VII 2012        --        TX
SHSUE 000173     1 VII 2007         --        NE
SHSUE 026343     20 VII 2013         M        FL
SHSUE 026340     19 IX 2013          M        OK
SHSUE 000195     2 IX 2007          --        KS
SHSUE 000162     2 VII 2007         --        KS
SHSUE 008488     12 VI 2013         --        LA
SHSUE 008519     17 VI 2013         --        LA
SHSUE 000692     1 VII 2011         --        TX
SHSUE 026347     31 VIII 2013        F        TX
SHSUE 026341     19 IX 2013          F        OK
SHSUE 026342     19 IX 2013          F        OK
SHSUE 008592     20 VII 2013        --        FL
SHSUE 008599     20 VII 2013        --        FL
SHSUE 008601     20 VII 2013        --        FL
SHSUE 008600     20 VII 2013        --        FL
SHSUE 008602     20 VII 2013        --        FL
SHSUE 008603     20 VII 2013        --        FL
SHSUE 008604     20 VII 2013        --        FL
SHSUE 008579     20 VII 2013        --        FL
SHSUE 008580     20 VII 2013        --        FL
SHSUE 008581     20 VII 2013        --        FL
SHSUE 008582     20 VII 2013        --        FL
SHSUE 008583     20 VII 2013        --        FL
SHSUE 008585     20 VII 2013        --        FL
SHSUE 008586     20 VII 2013        --        FL
SHSUE 014225     24 VII 2013        --        TX
SHSUE 014226     24 VII 2013        --        TX
SHSUE 026437     20 V 2014          --        TX
SHSUE 026438     20 V 2014          --        TX
SHSUE 007459     20 V 2014          --        TX
SHSUE 007460     20 V 2014          --        TX
SHSUE 007461     20 V 2014          --        TX
SHSUE 007462     20 V 2014          --        TX

Museum           County/Parish                 Locality
accession
numbers

USMN 00354813    N/A             C&O canal, Chain Bridge
USMN 00354821    Jackson         N/A
USMN 00354822    Jackson         N/A
IORI 00038262    Liberty         Torreya State Park
IORI             Holmes          Choctowhatchee River at US 90
IORI             n/a             Escambia River
IORI             Columbia        Santa Fe River
IORI             Clay            Black Creek, N. Prong, Hwy 209
IORI             Suwannee        Suwannee River, Suwannee State Park
IORI             Wakulla         St. Marks River at Newport on US 98
IORI             Gadsden         Near Apalachicola River, Hwy 90,
                                   Chattahoochee
IORI             Gadsden         Near Apalachicola River, Hwy 90,
                                   Chattahoochee
IORI             Jackson         Three Rivers State Park
IORI             Liberty         Ochlockonee River at Hwy 20
IORI             Washington      Choctowhatchee River at US 90
IORI             Jackson         Three Rivers State Park
IORI             Clay            Black Creek, N. Prong , Hwy 209
IORI             Columbia        Pond near Santa Fe River, Hwy 441
IORI             Alachua         Santa Fe River, near Hwy 235
USMN 00354826    Decatur         Spring Creek (mating pair)
GMNH             Morgan          N/A
GMNH             Clarke          N/A
GMNH             Sumter          N/A
GMNH             Clarke          N/A
GMNH             Sumter          N/A
GMNH             Clarke          N/A
GMNH             Clarke          N/A
GMNH             Clarke          N/A
GMNH             Clarke          N/A
GMNH             Clarke          1.1 mile SW of Winterville
GMNH             Clarke          N/A
GMNH             Fulton          Bolton
GMNH             Fulton/Dekalb   Atlanta
GMNH             Fulton          Bolton
GMNH             Putnam          Eatonton
GMNH             Whitfield       Dalton
GMNH             Decatur         Spring Creek
USMN 00354841    Lake            4-5 mi. N. of Olney
USMN 00354842    Lake            4-5 mi. N. of Olney
USMN 00355162    Tippecanoe      Wabash River, N. of Lafayette
USMN 00355164    Tippecanoe      Wabash River, N. of Lafayette
USMN 00354873    N/A             Iowa River; SUI Campus
USMN 00354875    N/A             Iowa River; SUI Campus
USMN 00354900    Sumner          Caldwell
USMN 00354901    Sumner          Caldwell
USMN 00354908    Green           Crailhope; Little Barren River
USMN 00354906    Green           Crailhope; Little Barren River
USMN 00354909    Madison         Eagle Lake
USMN 00354910    Madison         Eagle Lake
USMN 00354930    Montgomery      Barnesville; C&O canal
USMN 00354928    Montgomery      Barnesville; C&O canal
USMN 00354969    Washington      Stillwater
USMN 00354970    Washington      Stillwater
USMN 00354981    Oregon          N/A
USMN 00354979    Oregon          N/A
GMNH             Boone           N/A
USMN 00355192    Cherry          Along Niobrara R; Allen Bridge;
                                   4 mi. S of Sparks
USMN 00355192    Cherry          Along Niobrara R; Allen Bridge;
                                   4 mi. S of Sparks
USMN 00355200    Hunterdon       Lockatong Cr. On Hwy 29 N; Stockton
USMN 00355204    Hunterdon       Lockatong Cr. On Hwy 29 N; Stockton
USMN 00355209    Guadalupe       Santa Rosa
USMN 00355213    Swain/Jackson   Cherokee; Hiwassee River; Murphy
USMN 00355214    Swain/Jackson   Cherokee; Hiwassee River; Murphy
USMN 00355218    Erie            Huron R. ; 5 mi. S. of Huron
USMN 00355222    Erie            Huron R. ; 5 mi. S. of Huron
USMN 00355247    Paine           N/A
USMN 00355248    Garfield        N/A
USMN 00355254    York            Conewago Cr.; 5 mi. NW of Davidsburg
USMN 00355257    Greenville      Lakeside; a lake approx. 7 mi.
                                   S. of Greenville
USMN 00355256    Greenville      Lakeside; a lake approx. 7 mi.
                                   S. of Greenville
USMN 00355275    Campbell        Lafollette; Cove Lake
USMN 00355272    Davidson        Nashville; pool in Centennial Park
USMN 00355297    Cameron         Brownsville
USMN 00355303    San Jacinto     Near Coldspring
USMN 00355304    Wharton         El Campo
USMN 00355309    n/a             Dallas (mating pair)
USMN 00355306    San Jacinto     Near Coldspring
USMN 00391416    Fairfax         Quarry; Great Falls; 39.984722
                                   -77.250278
USMN 00391417    Fairfax         Swamp Trail; Great Falls; 38.984444
                                   -77..250556
USMN 00713680    Marshall        Dunkard; Fork Lake; 1.2 mi. S. of
                                   Majorsville off CR 15
USMN 00713676    Harrison        Good Hope; West Fork River
SHSUE 026337     Suwannee        Suwannee State Park along the river
SHSUE 026336     Suwannee        Suwannee State Park along the river
SHSUE 026339     Suwannee        Suwannee State Park along the river
SHSUE 026338     Suwannee        Suwannee State Park along the river
SHSUE 026335     Suwannee        Suwannee State Park along the river
SHSUE 026344     Suwannee        Suwannee State Park along the river
SHSUE 026345     Suwannee        Suwannee State Park along the river
SHSUE 026349     Taylor          Jim Ned Creek
SHSUE 026348     Taylor          Jim Ned Creek
SHSUE 026346     Taylor          Jim Ned Creek
SHSUE 000185     Tulsa           Mohawk State Park
SHSUE 000161     Brown           Yegua Creek
SHSUE 000189     Parker          Mineral Wells
SHSUE 001293     Walker          Raspberry Pond, Phelps, TX
SHSUE 000173     Omaha           Easley Creek
SHSUE 026343     Suwannee        Jim Ned Creek
SHSUE 026340     Waynoka         Waynoka Stream
SHSUE 000195     Sedewick        Pawnee Prairie
SHSUE 000162     Brown           Delaware River
SHSUE 008488     Calcacien       Sam Houston Jones State Park
SHSUE 008519     Washington      Bogue Chitto Stream
SHSUE 000692     Taylor          Jim Ned Creek
SHSUE 026347     Taylor          Jim Ned Creek
SHSUE 026341     Waynoka         Waynoka Stream
SHSUE 026342     Waynoka         Waynoka Stream
SHSUE 008592     Suwannee        Suwannee State River Park
SHSUE 008599     Suwannee        Suwannee State River Park
SHSUE 008601     Suwannee        Suwannee State River Park
SHSUE 008600     Suwannee        Suwannee State River Park
SHSUE 008602     Suwannee        Suwannee State River Park
SHSUE 008603     Suwannee        Suwannee State River Park
SHSUE 008604     Suwannee        Suwannee State River Park
SHSUE 008579     Suwannee        Suwannee State River Park
SHSUE 008580     Suwannee        Suwannee State River Park
SHSUE 008581     Suwannee        Suwannee State River Park
SHSUE 008582     Suwannee        Suwannee State River Park

SHSUE 008583     Suwannee        Suwannee State River Park
SHSUE 008585     Suwannee        Suwannee State River Park
SHSUE 008586     Suwannee        Suwannee State River Park
SHSUE 014225     Liberty         Big Thicket: Birdwatcher's Trail
SHSUE 014226     Liberty         Big Thicket: Birdwatcher's Trail
SHSUE 026437     Brewster        Rio Grande River
SHSUE 026438     Brewster        Rio Grande River
SHSUE 007459     Brewster        Rio Grande River
SHSUE 007460     Brewster        Rio Grande River
SHSUE 007461     Brewster        Rio Grande River
SHSUE 007462     Brewster        Rio Grande River

Table 2. List of Argia apicalis specimens processed
for molecular analysis.

Museum              Date       Sex   State
accession
numbers

SHSUE 026337    20 VII 2013     M     FL
SHSUE 026336    20 VII 2013     M     FL
SHSUE 026339    20 VII 2013     F     FL
SHSUE 026338    20 VII 2013     M     FL
SHSUE 026335    20 VII 2013     F     FL
SHSUE 026344    20 VII 2013     F     FL
SHSUE 026345    20 VII 2013     M     FL
SHSUE 026349    31 VIII 2013    F     TX
SHSUE 026348    31 VIII 2013    F     TX
SHSUE 026346    31 VIII 2013    M     TX
SHSUE 000185    1 IX 2007      --     OK
SHSUE 000161    23 VII 2007    --     TX
SHSUE 000189    10 VIII 2007   --     TX
SHSUE 001293    25 VII 2012    --     TX
SHSUE 000173    1 VII 2007     --     NE
SHSUE 026343    20 VII 2013     M     FL
SHSUE 026340    19 IX 2013      M     OK
SHSUE 000195    2 IX 2007      --     KS
SHSUE 000162    2 VII 2007     --     KS
SHSUE 008488    12 VI 2013     --     LA
SHSUE 008519    17 VI 2013     --     LA
SHSUE 000692    1 VII 2011     --     TX
SHSUE 026347    31 VIII 2013    F     TX
SHSUE 026341    19 IX 2013      F     OK
SHSUE 026342    19 IX 2013      F     OK
SHSUE 007459    20V2014        --     TX
SHSUE 007460    20V2014        --     TX
SHSUE 007461    20V2014        --     TX
SHSUE 007462    20V2014        --     TX

Museum          County/Parish             Locality
accession
numbers

SHSUE 026337    Suwannee        Suwannee State River Park
SHSUE 026336    Suwannee        Suwannee State River Park
SHSUE 026339    Suwannee        Suwannee State River Park
SHSUE 026338    Suwannee        Suwannee State River Park
SHSUE 026335    Suwannee        Suwannee State River Park
SHSUE 026344    Suwannee        Suwannee State River Park
SHSUE 026345    Suwannee        Suwannee State River Park
SHSUE 026349    Taylor          Jim Ned Creek
SHSUE 026348    Taylor          Jim Ned Creek
SHSUE 026346    Taylor          Jim Ned Creek
SHSUE 000185    Tulsa           Mohawk State Park
SHSUE 000161    Brown           Yegua Creek
SHSUE 000189    Parker          Mineral Wells
SHSUE 001293    Walker          Raspberry Pond, Phelps, TX
SHSUE 000173    Omaha           Easley Creek
SHSUE 026343    Suwannee        Jim Ned Creek
SHSUE 026340    Waynoka         Waynoka Stream
SHSUE 000195    Sedewick        Pawnee Prairie
SHSUE 000162    Brown           Delaware River
SHSUE 008488    Calcacien       Sam Houston Jones State Park
SHSUE 008519    Washington      Bogue Chitto Stream
SHSUE 000692    Taylor          Jim Ned Creek
SHSUE 026347    Taylor          Jim Ned Creek
SHSUE 026341    Waynoka         Waynoka Stream
SHSUE 026342    Waynoka         Waynoka Stream
SHSUE 007459    Brewster        Rio Grande River
SHSUE 007460    Brewster        Rio Grande River
SHSUE 007461    Brewster        Rio Grande River
SHSUE 007462    Brewster        Rio Grande River


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Please note: Some tables or figures were omitted from this article.
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Article Details
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Author:Sisson, Melissa S.; Santamaria, Carlos A.; Smith-Herron, Autumn J.; Cook, Tamara J.; Cook, Jerry L.
Publication:Florida Entomologist
Article Type:Report
Geographic Code:1USA
Date:Sep 1, 2016
Words:4712
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