A new species of Praticolella (Gastropoda: Polygyridae) from northeastern Mexico and revision of several species of this genus

2011·8.8 MB·English

by  Kathryn E Perez

.
THE NAUTILUS 125(3): 13-126, 201 Page 113
1 I
A new species of Praticolella (Gastropoda: Polygyridae) from
northeastern Mexico and revision of several species of this genus
Kathryn E. Perez
DepartmentofBiology
UniversityofWisconsinatLaCrosse
LaCrosse,WI54601 USA
[email protected]
ABSTRACT 1971; Neck, 1977; Hubrieht, 1984). The last taxonomic
revision of the United States species was by Pilsbry
A new species ofpolygyrid land snail ofthe genus Praticolella (1940) butexcludedthe fivenominal Mexicantaxa; these
from northeastern Mexico is described. This species has Mexican taxa were last revised by von Martens (1890-
established invasive populations in the United States and
1901).
CP.arbiebrblaenadnierainadna.haTshebeneenw cPorantfiucsoeledllawistpheciP.esgirsisseiomlialarantdo Praticolella griseola (Pfeiffer, 1841) is the only mem-
Pd.epgrriessesoelda,shbeultlwdiiftfherwshiitnebpeiignmgenltaartgeiro,n,haavfilnagttaenmedorweidreobbuosdty, borerinovfastihveegiennutshecuUrnrietnetdlyStcaotnessid(eUrSeAd)to(Dbuendienetr,od1u9c7e4d;
whorl, and a more oval-shaped aperture. The range of these Robinson, 1999). Individuals identified as P. griseola
two species does not appear to overlap with P. berlandieriana, have been recorded in the southeastern USA and the
which is restricted to central, north, and eastTexas. These two Caribbean since the 1920s. In addition, the United
species are also circumscribed and their taxonomic history is States Department of Agriculture (USDA) routinely
reviewed. This newtaxonomywas establishedusingmitochon- intercepts P griseola travelingwith shippingtrade.
drial 16S rDNAandcytochromeeoxidase subunit-I sequences This paper provides the means to distinguish several
aswellas geometric morphometricexamination ottheshellsot
morphologically similar species of Praticolella at least
eachspecies. ,
two ofwhich regularlytravel with shipments offruit and
Additional keywords: Mollusca, Invasive species, snail, mito- greenhouse plants. The data presented here will make
chondrial DNAanalysis,mitochondrial 16S rDNA,cytochrome evident that other Praticolella species also need taxo-
eoxidase, geometric morphometric analysis nomic attention, but this paper focuses on P. griseola
and the species which must be considered to sort out
the taxonomy ofthe invasive Praticolella species. These
include: P. griseola, P. berlandieriana (Moricand, 1833),
and P. strebeliana (Pilsbiy, 1899). In this study, I use 16S
INTRODUCTION rDNA (16S) and cytochrome c oxidase subunit I (COI)
mitochondrial DNA and shell geometric morphometric
Praticolella von Martens, 1892 is a genus of polygyrid analysis to provide the basic taxonomic and phyloge-
land snails found throughout the southeastern United netic information necessary for taxonomy, conservation,
States (USA), Mexico, and South to Central America and management of these invasive mollusks and native
(Pilsbry, 1940). Praticolella is currently composed of 15 congeners.
recognized species (Pilsbry, 1940; Hubrieht, 1984), nine
of which are species of conservation concern having
global heritage ranks of Gl, G2, or G3, indicating they MATERIALS AND METHODS
are considered critically imperiled, imperiled, orvulner-
DNA
able (Master, 1991; NatureServe, 2005). This paper aims Table 1 lists specimens examined for analysis, col-
to distinguish among several morphologically similar lection sites, latitude and longitude, and museum acces-
species ofland snail in the genus Praticolella sion numbers ofall specimens (also shown in Figure 1).
Species in Praticolella possess highly variable mor- Specimens examined in this study are deposited in the
phological characteristics often used in their taxonomy: AcademyofNatural Sciences of Philadelphia. Additional
the openness of the umbilicus; degree of depression specimens for morphometric analysis (also listed in
of the spire; and the pattern of pigmented bands on Table 1) were borrowed from the Florida Museum ol
the shells. This has ledto disagreements about taxonomy Natural History, Gainesville; American Museum of Nat-
of species in the genus (von Martens, 1890-1901; ural Histoiy, NewYork; Museum d'Histoire Naturelle,
Singley, 1893; Pilsbry, 1940; Cheatum and Fullington, Geneva; and Field Museum ofNatural Histoiy, Chicago.
Page ] 14 THE NAUTILUS, Vol. 125, No. 3
Table 1. Ijoealityinformation and museum number for specimens sequenced for DNA analysis followed by locality information
and museum number for additional lots included in morphometric analysis. In these additional lots all adult, complete shells were
photographed and included in morphometric analysis. ANSP numbers beginning with “A” represent lots preserved in alcohol.
Latitude and Longitude presented in decimal degrees.
Species Locality Museum Number Latitude Longitude
DNA
Localities for
specimens
Praticolellaberlandieriana 9km N New Braunfels, ComalCo. TX ANSPA22076, ANSP 29.7739 -98.1601
426024
Praticolellaberlandieriana 12.6km SE ofBlanco, Blanco Co. TX ANSPA22079 30.0204 -98.3300
"Praticolellagriseola” Point Isabellahigh school. PortIsabel, ANSPA22074, ANSP 26.077 -97.2271
Cameron Co CameronCo. TX 426021
Praticolellagriseola 9.5km N ofPuenteArroyo Hondo, ANSPA22080 20.0883 -96.9106
(N ofVeracruz),VC, MEX
Praticolellagriseola 9km N ofPapantla,VC, MEX ANSPA22082 20.46389 -97.34111
Praticolellagriseola Tula, NL, MEX ANSPA22081 23 -99.7167
Praticolellagriseola San Rafael,VC, MEX ANSPA22075, ANSP 20.2 -96.85
426022
Praticolellagriseola Veracruz,VC, MEX ANSPA22078, ANSP 19.2147 -96.16028
426029
Praticolellagriseola La Mancha,VC, MEX ANSPA22073, ANSP 19.6061 -96.39889
426019
Praticolellagriseola 1.5 mi S ofJimenez, TMP, MEX ANSP426028 24.2167 -98.4667
Praticolellagriseola 1 km E Howey Height, Lake Co. Fl ANSPA22084 28.667676 -81.746159
Praticolella mexicana Diente, ~5km S of Monterrey, ANSPA22072ANSP426018 25.47728 -100.28673
NL, MEX
Praticolella mexicana Anahuac, NL, MEX ANSPA22086 27.2519 -100.125
PPrraattiiccoolleellllaa mmeexxiiccaannaa ~Pu1e0ntkemSWanoRfoMdornitgoe,rrCeyO,AHNL,,MMEEXX AANNSSPPAA2222007879, ANSP 2285..666768861 --110000..49418217
426025
Praticolella mexicana 3km N Agua Buena, RioTamasopo, ANSPA22087 21.9419 -99.395
MEX
SLP,
Praticolella mexicana Key Largo FL ANSPA22091 25.17202 -80.3666
Praticolella mexicana 2km N Agua Buena, SLP, MEX ANSPA22088 21.96426 -99.38899
Praticolella mexicana Hidalgo, HidalgoCo. TX ANSP426020 26.1 -98.263
Praticolella mexicana 15km SWof Linares,where road ANSPA22101, ANSP 24.7573 -99.6581
to CajaPintaruns alongstream, 426032 (Paratypes)
NL, MEX
Praticolella mexicana ~5km E CiudadVictoria, TMP, MEX ANSPA22085 23.7333 -99.1333
Praticolella mexicana 20km N Mante,TMP, MEX ANSP426027 22.8833 -99.0167
Praticolella mexicana -int USDA - JFKinternational airport ANSPA22094 40.6501 -73.9496
interception from Dominican
Republic
Praticolella mexicana - int USDA - Progresso from Mexico ANSPA22093 26.0923 -97.9572
Praticolella mexicana Hell, GrandCayman USDA-APHIS Collection 19.381774 -81.415936
Praticolella mexicana Staniard Creek, Andros Island, ANSPA22090 24.844126 -77.892870
Bahamas
Morphometries localities
P. mexicana 15km SWofLinares. Ingrass along ANSP426031 (Holotype), 24.7573 -99.6581
fence at S endofparknext ANSP426032,ANSP
basketball court, nexttosmall A22101 (Paratypes)
stream, where Mx58toCajaPinta
runs alongstream, NL, MEX
Canoas, SLP Inparkin town on road ANSP426026 21.94405 21.94405
through town bystream.
P. berlandieriana Milam Countynear Brazos River, FMNI4 259146 30.8289 -96.6706
4.7 mi NE ofGause, USA, Texas
3.3 mi N New Braunfels, Comal Co. ANSP426023 29.7635 -98.1547
20 M from Guadalupe Riveracross ANSP426024ANSPA22076 29.7739 -98.1601
from Guadalupe Canoeing9km N
New Braunfels, ComalCo. TX
MHNG
“Habitele Mexique, dans laprovince 37027
deTexas.”
Continued
( )
1
K.E. Perez, 2011 Page 115
Table 1. (Continued)
Species Locality Museum Number Latitude Longitude
“Praticolellagriseola” Cameron Count)', Port Brownsville, FMNPI 259156 25.949904 -97.402715
Cameron Co TX, USA
0.3mi E Point Isabellahigh ANSP426021 ANSPA22074 26.077 -97.2271
school. Port Isabel, Cameron
Co. TX
FMNH
Praticolellagriseola Nicaragua 33486 12.865416 -85.207229
Sequences were assembled in Sequencher™ 4.0.5
(Gene Codes Corporation, Ann Arbor, MI) or Geneious
Pro 3.5.6 (Drummond et al., 2006) and aligned in Muscle
(Edgar, 2004). Garli 0.951 (Zwickl, 2006) was used to
carry out maximum likelihood (ML) estimation of rela-
ML
tionships. The analysis was carried out using Garlis
default settings, as was an additional 100 replicate boot-
strap analysis.
Shell Morphometric Analysis: In total, 237 spec-
imens representingthe followingfivespecieswereexam-
ined: P. berlandieriana (46 specimens), “P. griseola"
Cameron County (32), P. gri.seola (68), Praticolella new
species (described below) (85), P. strebeliana (5) (Speci-
mens used for DNA are listed in Table 1). We had diffi-
culty in identifying to species specimens without living
tissue for sequencing (shell-only specimens). Therefore,
Figure 1. Map of specimens used for DNA analysis and only a limited number of individuals from museum col-
additional localities of specimens used for morphometric lections could be used for morphometric analysis, in
analysis. addition to the individuals for which we gathered
sequence data for morphometric analysis; this number
mostly includes additional individuals or shell-only
Individual sequences are available on Genbank, 16S: collections from the same locality as individuals with
DQ085935-DQ086020, COI: DQ086021-DQ086095. sequenced DNA. Color images were captured with a
Outgroups were included from other members of the tripod-mounted. Canon PowerShot S3IS digital camera.
Polygyrini (Emberton, 1995), specifically, Polygyra Twenty-five landmarks (Figure 2) were digitized using
cereolus (Miihlfeld, 1816) and Polygyra septemvolva tpsDig 1.31 (Rohlf, 2001).
(Say, 1818). Geometric morphometries analyses were carried out
using the Integrated Morphometries Package, this
includes the programs CoordGen, PCAGen, CVAGen,
Molecular Data Analysis: Total genomic DNA was TwoGroup, and Regress 6 listed below (IMP software
extracted from several milligrams oftissue digested with suite; Sheets, 2003) were used to examine shape varia-
CTAB lysis buffer and purified through standard phenol- tion through principal component analysis (PCA) and
chloroformprocedures (Palumbietal., 1991). Degenerate canonical variance analysis (CVA). PCA is a technique
primers (5'-YRMCTGTTTAWCAAAAACAK-3' and 5'- for simplifying descriptions ofvariation among individ-
CCGGTCTGAATCCAGATCABGT-3') were designed uals, while CVA simplifies descriptions of differences
from the Palumbi et al. (1991) primer pair and Genbank between pre-determined groups (Zelditeh et al., 2004).
sequencesthatamplifieda~450bp fragmentofthe mito- In traditional morphometric analysis, PCA generallysuf-
chondrial 16S gene. The Folmer et al. (1994) primers fers from the overwhelming influence ofsize across the
were used to amplify a ~600 bp fragment of the mito- newly generated axes. However, geometric morphomet-
chondrial cytochrome oxidase c subunit I gene (COI) ric analysis eliminates size as a factor, yielding examina-
gene. Fragments were amplified by PCR, purified tions of shape solely. In PCA, no a priori assumptions
through gel extraction, and sequenced using BigDye 3. are needed to group individuals. In contrast, CVA
chemistry on an ABI 3100 automatic genetic analyzer. determines the set of axes that best discriminates
Specific protocols for amplification and sequencing can between groups; therefore an a priori assumption
be found in Perezetal. (2005). of group membership is necessary. For CVA analysis,
=
Page 116 THE NAUTILUS, Vol. 125, No. 3
Llorida) or United States Department of Agriculture
(USDA) interceptions and were initially identified as
P. griseola, which is well known as invasive. The DNA
tree also shows monophyletic lineages from both near
Victoria, Tamaulipas, and near Mante, Tamaulipas.
These are herein considered part of Praticolella new
species , but further work is needed to examine popula-
tion level versus species level differences among these
lineages.
IndividualsconformingtothemoiphologyofP. griseola,
including topotypic material, form a clade (100% boot-
strap support; labeled P. griseola on Ligure 3) that has
deep subdivisions between different populations in
DNA
the mitochondrial and includes an individual from
an invasive population in Lake County, Florida. The
Figure2. Praticolellamexicanaindividual(#436) from 15km P. griseola clade is resolved as sister (75% bootstrap
SWofLinares, Nuevo Leon, (ANSP 426031) Mexico showing support) toamonophyleticgroupol individuals from Soto
landmarks used in the morphometric analysis. This individual la Marina, Tamaulipas. Individuals from asmall, morphol-
usedinboth DNAandmorphometricanalyses. ogically unique, disjunct population of P. griseola were
sampled in this study and are called here: “P. griseola”
Cameron Co. (Pilsbry, 1940; Rehder, 1966). This popula-
individuals were grouped according to the clades identi- tdiuoen ftoorpmoeodrasudpisptoirntcticnltaidnesspeopratriaotneoffrtohmeP.trgerei,seroellaat,ibount-
fiedbythe molecular analysis. ships remainuncertain.
Landmarkcoordinates were importedintoCoordGenGl Individuals from as close to the type locality of
ParnodcrcuosntveesrtseudpetroimPprooscirtuisotnesmedtihstoadnsc.esAuMsiAngNOleVasAt scqaurarrieesd wP.itbheirnlan3c0liekrima)nafasorcmoulad bcleaddeetesrismtienredto(deottahielrs bTeelxoaws,
out in SYSTAT 8.0 was used to examine differences Praticolella species (84% bootstrap), including individ-
in shape between species. Pairwise comparisons between uals ofP. trimatris Hubrieht, 1983, P. pachyloma (Menke
all populations were performed in TwoGroup6c with in Pfeiffer, 1847) and P. taeniata Pilsbry, 1940.
Bonferroni correction to determine if there were sig-
nificant shape differences. A PCA was performed with Shell Morphometric Analysis: Differences in shell
PCAGenGg on die data with a posteriori groups assigned shape in Praticolella were assessedusing GM (Figure 3).
by locality. A CVA was also performed usinDgNCAVAGen6h Shell variation is traditionally quantified through
with groups defined bythe clades from the analysis. straight-line shell measurements and ratios and used to
The difference in shape between each species was exam- distinguish between individuals and populations at the
ined directiyusingRegress6. species level (e.g., Heller et ah, 2005; Tanaka and Maia,
GM
Abbreviations and Text Conventions: MHNG = 2006). Recently, hasbeen employed in examinations
Museum d’Histoire Naturelle, Geneva; LMNH = Lield ofsnail shells, both toprovide direct size-free analyses of
Museum ofNatural History; ANSP =AcademyofNatu- shellshapeandtoanswerbroaderevolutionaryquestions
(Pfenninger and Magnin, 2001; Conde-Padin et ah,
ral Sciences of Philadelphia (ANSP numbers beginning
wUintihte“dA”StraetpersesDeenptarlottmsepnrtesoefrvAegdriicnulatlucroeh;ol)G;MUS=DAgeo=- 200T7h;eHfaiyrsetsPeCtAaha,xi2s00(7P)C.I) explained 31.5% ofthe vari-
vpmaeortnireainnccte.maonLraapltyhisoitmsu;edetrMaiAncdNaOLnaoVlnAygs=iist;udmPuelCtpAirveas=reinapttreeidncaiinnpaladylescicsiommao-lf as(tiPigCon3ni),f,itcah1ne3t.s1de%ic.fofnAedreM(nAcPeCN2aO)mVeoxAnpgloanignrteohduep1sP5C.(3HA%o,tsecalolnridensgt-hfLeoautwnhldieryda
degrees. Trace=2.012, F-Statistic=30.669, df=15, 686, p<0.000).
Pair-wise comparisons of all species assessed by
Goodall’s F test showed that snails from each species
RESULTS had significantly different (p<0.01) shapes.
Each species was comparedpairwise usingTwoGroup
Molecular Phylogeny: Maximum likelihood analysis to carry out Goodall’s F-test. Pairwise comparisons were
of417 bp of 16S and 493 bp ofCOI yielded asingle tree followed by a Bonferroni correction. This analysis found
(Ligure 3). Praticolella new species (to be described that each ofthe species’ means are significantly different
below) comprised a well-supported monophyletic elade. (p<0.001 in allcases). Thelectotypes (ANSP411457and
Many individuals included in this clade were initially 77128) ofPraticolella strebeliana are distinct on the first
identified as P. berlanclieriana due to their geographic three PC axes (Figure 4) with the highest difference in
location in northeastern Mexico. Some individuals inthis mean value from the other species (Distance in mean
clade were from invasive populations (Bahamas and value from P. griseola=0.0967; P. berlanclieriana 0.0916;
7
1
K.E. Perez, 201 Page 1 1
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64 • P.traaRiPsp.aai2p^.tfeRreiSawtiSarRrd<aiFpisiarta,TteXeiB1^5TT!VXP1143*2 P. berlandtenana
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Figure 3. Molecular phylogeny of Praticolella. Maximum likelihood phylogram based on 16S and COI mitochondrial DNA.
The species discussed in this paper are marked by grey boxes. Numbers on branches are ML bootstrap values. Outgroups not
shown.
Page 118 THE NAUTILUS, Vol. 125, No. 3
-o.l -0.05 0 0.05 0.1 -0.07 -0.02 0.03 0.08
Figure 4. Principal component axes 1, 2, and 3. Percentages indicate the percent ofthe variance explained by each axis. Large
symbols represent group means: Praticolella mexicana (•); P. berlandieriana (0); P. griseola (); “P. griseola” Cameron Co (A)
P. strebeliana (+).
-0.01 0 0.01 -0.01 0.00
Figure 5. Canonical axes 1, 2, and 3. Discriminant scores calculated to maximally separate the tive species. Left: the first and
secondcanonicalaxes. Right: thesecondandthirdcanonicalaxes. Largesymbols representgroupcentroid. Praticolella mexicana (•);
P. berlandieriana (0); P. griseola (); "P. griseola”Cameron Co (A) P. strebeliana (+).
Praticolella new species=0.0723). Although the species nificantly different from each other (Figure 5). The
are significantlydifferent, visual examination ofFigure 4 resultingplot ofCV 1 and CV2 shows very little overlap
shows there is a great deal of overlap in the shape among species, although a few individuals ofP. griseola
variation present in each species. Praticolella new spe- (three of64) overlap into the new species’ shape space.
cies is the most distinct in shape (Distance in mean One individual of Praticolella new species grouped with
value from P. griseola=0.0448; P. berlandieriana=0.0590; P. griseola. The plot of CV2 and CV 3 widely separate
P. strebeliana=0.0723), but Praticolella new species, P. strebeliana and P. griseola from the other species.
P. griseola and P. berlandieriana also have a great deal Finally, “P. griseola” Cameron Co. is significantly differ-
of overlap in shape (Figure 4; Distance in mean ent from the other species and distinct from P. griseola
value=0.0518). CVA of each population yielded four and Praticolella new species in all analyses. Figure 6
distinct axes (p< 0.05) where all centroids were sig- shows how shell shape differs from Praticolella new
K.E. Perez, 2011 Page 119
Figure 6. Change in shellshapebetween species. Lett: Shape changeto P. berlandieriana fromP. mexicana. Right: shape change
toP. griseola from P. mexicana. Shape changeis exaggerated3Xbyvectorarrows toeaseinterpretation.
species to P. berlandieriana and (left) and from nine complete and incomplete bands, radiating lines of
Praticolella newspecies to P. griseola (right). white pigment, and streaks of white pigment on body
whorl. Umbilical whorls with line growth lines but no
spiral striae and usually brown/gray colored with no
SYSTEMATICS
white pigment, shiny. Average shell height=7.57,
width=10.87, umbilicus width=0.69 mm, 5-5.6 whorls
Family Polygyridae Pilsbry, 1930
(Table 2).
Genus Praticolella von Martens, 1892 Type Material: Ilolotype ANSP 426031, 27 May
1992, Ned E. Strenth (Figures 7-10). Paratypes ANSP
DorcasiaBinney, 1878: 356. 426032 andalcohol-preservedspecimens ANSPA22101.
Praticola Strebel and Pfeiffer, 1880: 38.
Other material examined: Hidalgo, Hidalgo Co. TX, 21
Praticolella von Martens, 1892: 138. Sept. 1991, Ned E. Strenth (Figure 11), ANSP 426020;
College of the Bahamas Research Station, Staniard
Type Species: Praticolella ampla (Pfeiffer, 1866), by Creek, Andros Island, Bahamas, 20 May 2005, K. E.
original designation. Perez, ANSP A22090 (Figure 12); Canoas, San Luis
Potosf, Mexico, 23 July 2002, K.E. Perez. J.B. Pollock,
Diagnosis: Shell small, globose to slightly depressed ANSP 426026 (Figure 13).
with a conic spire of4.5 to 5.75 whorls. Aperture either Type Locality: 15 km SW of Linares, Nuevo Leon,
slightly or greatly reflected and without denticles. Nar- MX
Mexico, in grass next to small stream where 58
rowly umbilieate (Pilsbry, 1940). Penial diverticulum
to Caja Pinta runs along stream, 24.757331 N,
long and at least twice the volume of the penis -99.658111 W.
(Emberton, 1995). Bifurcate or trifurcate penial retrac-
tor muscle (Emberton, 1995). Distribution and Habitat: Widely distributed in
northeastern Mexico and southTexas on the eastern side
Distribution: United States: Florida, Georgia, Ala- of the Sierra Madre Oriental. There may be native
bama, Mississippi, Louisiana, North Carolina, Texas, populations in south Texas, though the collections exam-
Mexico south to Panama, Caribbean islands. ined were all from disturbed habitat or greenhouses.
United States Department ofAgriculture (USDA) often
Praticolella mexicana newspecies intercepts this species attheTexas/Mexicoborder. Intro-
duced populations were found in Florida, Bahamas,
Figures (7-13)
Grand Cayman Island, Dominican Republic, Haiti, and
Praticolella berlandieriana (Moricand, 1833): Fischer and Cuba. Several USDA interceptions were from Jamaica
Crosse, 1872: 256-257;Vanatta, 1915: 194, fig. 1 (genitalia); and Turkey. The native range of this species is most
Pilsbry, 1940: 695, fig. 427a; Rehder, 1966: 290-291, fig. 20; likely northeastern Mexico, north and east ofthe Sierra
Correa-Sandoval, 1993: 685; Correa-Sandoval, 1996: 137; Madre Oriental. First, the basal lineages in this clade
Correa-Sandoval, 1999: 15; Correa-Sandoval, 2000: 493; are all found in this region of Mexico. Second, collec-
Correa-SandovalandCastro, 2002: 238. tions from this region of Mexico predate the collection
Praticolellagriseola Pfeiffer, 1841: Robinson, 1999: 415. of this species in the Caribbean or Florida. This species
was first reported as introductions collected in dis-
Description: Shell umbilieate, globose to somewhat turbed habitats of Florida in the early part of the
depressed-globose. Lip thin, reflected to slightly cover 1900’s.
umbilicus. Banding extremely variable, ranging from Thepreferreddietofthisspeciesisunknown;however,
unbanded brown shell, unbanded white shell, to havinOg ithasbeen found onornamental (greenhouse) plants and
7
.
Page 120 THE NAUTILUS, Vol. 125, No. 3
Figures 7-13. Shells ol Praticolella mexicana new species. 7-10. Holotype, SWofLinares, Nuevo Leon, Mexico, 27 May 1992,
Ned E. Strenth, ANSP426031; side, top, andbasalviews ofthe shell andembryonicwhorls, w=9.58 mm, h=6.59 mm, 5.25whorls.
11. ANSP426020, Hidalgo, HidalgoCo.Texas,w=10.59,h=7.46,5.25whorls,21 Sept. 1991, NedE. Strenthcoll. 12. ANSPA22090,
College ofthe Bahamas Research Station, Staniard Creek, Andros Island, Bahamas, w=10.57, h=7.63, 5.25 whorls, 20 May 2005,
K.E. Perezcoll. 13. ANSP426026, Canoas, SanLuis Potosi, Mexico,w=9.79mm, h=7.1Smm,5.25whorls,23July2002, K.E. Perez,
J.B. Pollockcolls.
Table 2. Shell measurements forthe threespeciesofPraticolella Onlyadultshellswith afulllipweremeasured: P. berlandieriana
(n=24), P. griseola (n=36), P. mexicana (n=37). Valuespresent, from top, range, mean andstandarddeviation. Abbreviations: h: shell
height; w: shell width; aph: aperture height; apw: aperture width; umb: umbilicus width; # of whorls - number ofwhorls.
Species h (mm) w(mm) aph (mm) apw(mm) umb (mm) #ofwhorls
P. mexicana 6.3-9.19 9.38-12.27 4-6.77 4.85-6.92 0.4-1.08 5-5.6
7,57±0.61 10.87i0.78 5.28i0.51 5.97i0.60 0.69i0.15 5.30i0.17
P. griseola 8.32-11.29 5.8-7.92 4.4-6. 4.16-5.75 0.38-1.03 4.75-5.5
9.65T0.78 6.91i0.51 5.34i0.47 4.88i0,39 0.71i0.16 5.12i0.16
P. berlandieriana 9.92-11.73 7.34-8.75 4,5-6.14 4.65-5.77 0.64-1.18 5-5.5
10.49i0.49 8.16i0.40 5.52i0.42 5.19i0.29 0.85i0.13 5.35i0.22
is commoninsugarcane, citrus, mango,banana, aloe, and snails possess many of the typical characteristics of
papaya plantations (USDA interception records). The invasive snail species, such as living at high population
USDA has intercepted this species on shipments of densities in shrubs, tall grass, and under trash. This spe-
mangos,papayas,ornamentalplants,andfurniture.These cies is often foundclimbingwalls andgrass.
K.E. Perez, 2011 Page 121
Etymology: Named in reference to the native distri- Helix (Praticola)griseola (Pilsbry, 1891): 313.
bution ofthis species in Mexico. Praticolellagriseola Pilsbry, 1940: 690-692, tig. 425.
Taxonomic Remarks: Praticolella mexicana is similar Description: Praticolella with a robust, umbilicate,
to P. griseola, but differs in being larger, having a more depressed-globose shell. Number of pigmented bands
robust, depressed shell with white pigmentation, a flat- on bodywhorl ranges from 1 to 8with mostshells having
tened wide body whorl, and a more oval-shaped aper- 2 or 3. Most individuals possess a complete cinnamon
ture. Figure 6 (right) illustrates the difference in shape colored mid-bodywhorl band. Aperture lunate to round
between P. mexicana and P. griseola. The bodywhorl of with a thin reflexed lip. Shell obliquely striate. Average
Praticolella griseola (Figure 14) is more rounded with a shell height=9.65, width=6.91, umbilicus width=0.71
rounded aperture. The insertion of the peristome is mm, 4.75-5.5whorls (Table 2).
much closer to vertical in P. griseola and horizontal in
hfP.eraomvmeixeirPc.asnhmaee.lxliPcraaantndiacowlbielydlealhybaeverilxnagpnadnaideertdialalnelrai,pis(mFduiicgshutrientghu6iicsklheeerfd,t TraylpHeisMtaotryerMiuals:eumSyonftyLpoens,do6ni2nd0i1v1i0d1u7al9s.,FMiegxuirceos.1N4a-t1u9-.
and 20).
Praticolella strebeliana was included in this study as Distribution and Habitat: Pfeiffer (1841) gave the
it was described from Diente Mine near Monterrey, type locality of P. griseola as Veracruz. This species is
Nuevo Leon Mexico (Pilsbry, 1899) within the range ol native toVeracruz and southernTamaulipas and has also
collections of P. mexicana. This species was described been introduced to South Florida and New Orleans,
as completely brown with no bands; however, because Louisiana. Due to restriction of populations in the
occasionally P. mexicana individuals have no bands Yucatan and Guatemala to disturbed areas, it is consid-
(populations from Hidalgo, Texas are all bandless with ered invasive there as well (Harry, 1950). However,
a brown base color), I thought it necessary to con- native populations in Guatemala have not been ruled
sider P. strebeliana as potentially having priority before out by this data. The complete range ofthis species will
naming this new taxon (P. mexicana). Therefore, to test need further work to be hilly circumscribed. Specimens
whether P. strebeliana was an appropriate name, labeled P. griseola in museum collections are often
Praticolella specimens from Diente, the type locality of P. mexicana.
P. strebeliana, were collected for both DNA and morpho-
metric analyses. Several collecting attempts at the type Taxonomic Remarks: Praticolella griseola was de-
locality yielded no unbanded shells fitting the descrip- scribed by Pfeiffer (1841) in a short paragraph without
tion ofP. strebeliana. All individuals sequenced from ibis illustration. The specimens are attributed to Hegewisch,
localityarewithinthe P. mexicana clade. Therefore, I have referring to the physician and botanical collector
no DNA evidence to distinguish P. strebeliana. However, Dr. Ernst Friedrich Adoph Ilegewisch, who lived in
morphometric analysis of the type specimens (ANSP Oaxaca, Mexico, around 1836-1840 (Pritzel, 1864).
77128 and 411457) of P. strebeliana showed that they Pfeiffersprimarycollection (collection532)was lostwith
were veiy distinct from P mexicana, P. berlandieriana, the destruction of the Stettin Museum (Dance, 1986).
and P. griseola (Figures 4 and 5). Praticolella strebeliana However, some Pfeiffer material resides in the Natural
has a frosted, corneous shell that is more globose than History Museum in London (NHMUK) including a lot
P. mexicana, as well as a downward tilted lip and greater of 6 specimens labeled H. griseola Mexico Pfr in
degree ofcontraction behind the lip. Based on the differ- Pfeiffer’s handwriting (handwriting identified by Jona-
ences in shell morphology and morphometric analysis than Ablett, Curator of Non-Marine Mollusca and
I am considering P. strebeliana distinct from P. mexicana. Cephalopoda, NMHUK, pers. comm.). These specimens
In texture and coloration ol the shell, P. strebeliana is were also labeled “M.C.” indicating they came from the
more similar to P. flavescens than the other Texas or Hugh Cuming collection. While it is notpossible to con-
Mexican taxa. clude that these specimens were from the original type
series, Pfeiffer’s handwriting on the label indicates they
Praticolella griseola (Pfeiffer, 1841)
are probable syntypes.
Helix cicercula Ferrusae in collection = griseola according to In the phylogenetic tree (Figure 3), topotypic spec-
Pfeiffer 1848, 1: 337. imens conforming to the original description of
Bradi/baenapisam Beck, 1837: 18 (nomen nudum.) P. griseola formed a monophyletic lineage with individ-
Helixgriseola Pfeiffer, 1841: 41; Pfeiffer, 1848: 337. uals from an introduced population in Florida as well as
Helixalbocincta Binney, 1851: 109, 128. specimens from the coastal plain ofVeracruz and north
Helixalbo-zonata Binney, 1857: pi. 49, tig. 2. into Tamaulipas.
HHeelliixxsalpbloelnidniedautlaaGAonutlodn,in18B3i9n:ne3y6,. 1n85o7m:en34.nudum). Praticolella griseola has been the subject of much
( taxonomic contention. Von Martens (1890-1901) and
Dorcasia griseola Pleilfer, 1841; Binney, 1878: 348, tig. 231
(jaw), pi. vii, fig. v (teeth). Singley (1893) statedthat II. griseola and H. (Praticolella)
Helix berlandieriana var. griseola Pfeiffer, 1841; von Martens, berlandieriana are connected by many intermediate
1892: 140, pi. 7, figs. 15-17. forms and cannot be maintained as distinct species.
Page 122 THE NAUTILUS, Vol. 125, No. 3
Figures 14-19. Shells of Praticolella griseola. 14-19. Syntypes, Mexico, Pfeiffer material, H. Cuming Collection MNHUK
20110179. 14-17. Side, top, and basal views ol the shell and embryonic whorls. Scale bar — 1 mm (Figure 14). 18. Side view of
additionalshell from same lot. 19. Sideviewofadditionalshell from same lot.
However, Pilsbry (1940) found no connecting links variety of habitats and climates (Rehder, 1966; Neck,
between P. griseola andP. berlandieriana andfurtherpro- 1977). Rehder's (1966) “races” of P. griseola included
,
posed that they formed an ecological pairwith P. griseola the populations around Veracruz, Mexico and a second
living in warmer more humid regions, and P. race comprisedof asmall, unique, disjunctpopulation in
berlandieriana living in cooler, semiarid country. How- Cameron County in south Texas. Individuals from this
ever, Cheatum and Fullington (1971) stated, without “race” were sampled in this study and are referenced
presentingevidence, that, tine to interbreeding, in alarge herein as “P. griseola" Cameron Co. (Pilsbry, 1940;
assortment ofshells representing all species (meaning all Rehder, 1966). These snails have a thinner lip and a
species present in south Texas) from the same geographic dark-colored basal whorl. Taxonomic placement of this
area it is difficult to determine where one species ends population is outside of the sampling and scope of this
and anotherbegins. study, but the mitochondrial DNA results indicate thatit
Praticolellagriseola has been suggestedto be made up is distinct from P. griseola and from other nearby
of a number of well-characterized “races” living in a Praticolella species and remains to be described.