Volume 132, Numbers 3-4 December 21, 2018 ISSN 0028-1344
A quarterly devoted to malacology.
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José H. Leal
The Bailey-Matthews National Shell Museum
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M. G. Harasewych Department of Invertebrate Zoology National Museum of
Natural History Smithsonian Institution Washington, DC 20560 USA
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North Carolina State Museum of Natural Sciences
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Gary Rosenberg
Department of Mollusks
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CONTENTS
M.G. Harasewych Manuel J. Tenorio
Bianca Campagnari Daniel L. Geiger
Richard L. Squires
Daniel Abbate Luiz Ricardo L. Simone Daniel C. Cavallari
Shugian Zhang Peng Wei
Kazutaka Amano Krzysztof Hryniewicz Robert G. Jenkins
Omar Mejia Benjamin Lopez José Ma. Reyes-Gomez
Author Index ..........................
Meo riliuUs
Volume 132, Numbers 3-4 December 21, 2018 ISSN 0028-1344
The genus Cerion (Gastropoda: Pulmonata: Cerionidae) on San Salvador [Watling Island], Bahamas: A geometric morphometric analysis of shell TAA ONT] OL OK CH COVEY tsb acn eR heat Ao EUROS. SOR ba OE Rae RED En rr ee al
How many micromollusks are there? A case study on species richness in Hawai'i, with the description of a new species of Murdochella (Gastropoda:
IOULOOTIICEYS)) sancneretnauhaceernnadetee ae aannene cae RE senet Netee cieheicl Ee cect ochre ato etecateceecere eae §3 Catalog of the taxonomic updates of northeastern Pacific Late Cretaceous shallow-marine bivalves and gastropods named from 1874 to 1966 ............0.... 9] Anatomy of Engoniophos unicinctus from Isla Margarita, Venezuela
(Gastropoda: Caenogastropoda: Nassariidae), with a discussion on the lowmrcctinioimacsavaticl 1ElEVSIOINGLOTS) .0..2.ssqcoscenneaceeosecvossosncdx0s90aqJo00900s00Ke0e7en0G0badBENEAODDHOG 10]
Mericella zhangsupingae, a new cancellariid species from the South China Sen (Castropocles CancalletttC®)) ccosasadonscoscenocobancscdadabéodeasodandadeanoncbosesacadosacbHGHORE 113
A newly discovered Paleocene species of Boreocomitas (Gastropoda: Pseudomelatomidae) from eastern Hokkaido, Japan, with implications for thesbiogeourapinyazolstnesealeocencyb eninge Stal tems eee sean en 117
Three new species of the genus Humboldtiana (Gastropoda: Pulmonata: Helio @ | Gltsirrni Glave) efi @ Tn x © eee 124
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THE NAUTILUS 132(3-4):71-82, 2018
Page 7]
The genus Cerion (Gastropoda: Pulmonata: Cerionidae) on San
Salvador [Watling Island], analysis of shell morphology
M. G. Harasewych
Department of Invertebrate Zoology, MRC-163 National Museum of Natural History Smithsonian Institution
PO Box 37012
Washington, D.C. 20013-7012, USA
Bahamas: A geometric morphometric
Manuel J. Tenorio
Departamento de CMIM y Quimica Inorganica-IN BIO Facultad de Ciencias, Torre Norte, 1“ Planta Universidad de 11510 Puerto Real Cadiz, SPAIN
Cadiz
ABSTRACT
Morphometric analyses of shell shape of living specimens of
Cerion inhabiting San Salvador Island segregate samples into two primary phenotypes, one inhabiting the westem and
southem coasts of the island, the other the eastern and much of
the northem coast. These are concordant with phenotypes identified in prior morphometric studies. Lectotypes are designated for Cerion watlingense Dall, 1905; C. incon- spicuum Dall, 1905; C. inconspicuum lacunorum Dall,
1905; and C. coloni Bartsch, 1924. The lectotype of
Cerion watlingense Dall, 1905 falls within the western and southern phenotype, and is the oldest name available for this taxon. The lectotype of Cerion coloni Bartsch, 1924, a validly introduced, but previously unrecognized taxon, and the holotype of Cerion rodrigoi Gould, 1997 both fall within the group containing the east coast populations, with Cerion coloni Bartsch, 1924 being the oldest available name for this pheno- type. A third, previously unrecognized phenotype, represented by a single inland population, is morphologically similar to the lectotype of Cerion inconspicuum Dall, 1905, which is the oldest available name for this phenotype. The geographical distribution and chronological succession of these phenotypes since the late Pleistocene is reviewed in the context of both the single and multiple colonization models for the arrival of Cerion on San Salvador, and the evolutionary and taxonomic corollaries of each model are discussed.
INTRODUCTION
Nearly all modern papers dealing with land snails of the
genus Cerion are prefaced by an acknowledgment of
ihe extreme morphological diversity among the many, small populations of these snails. This is usually followed by a lamentation (e.g., Woodruff, 1978) about the pro- fusion of names |ca. 600] that have be en applied to the various phenotypes, mostly in the late 19"? and early 20" centuries. Nearly all of these named taxa are allopatric, but many interbreed freely when brought into contact (see e.g., Bartsch, 1920; Woodruff and Gould, 1987).
Throughout the range of Cerion (from the Keys and barrier islands of tropical Florida, throughout the Baha- mas, Greater Antilles, Cayman Islands, western Virgin Islands, and the Dutch Antilles, but absent from Jamaica, the Lesser Antilles, and coastal Central and South America), there are but three reported cases in which two different phenotypes co-occur sympatrically, one in Cuba (Mayr, 1963), and two on Great Inagua (Gould and Woodruff, 1990: Goodfriend and Gonitel 1996).
Earliest records of Cerionidae date from the Creta- ceous of Montana (Roth and Hartman, 1998), yet the overwhelming majority of taxa (>98%) are from late Pleistocene, ieislbcene. and especially Recent faunas, particularly those of Cuba (29%) and the Great Bahama
Bank (51%) (Har asewych, 2012: Table 1). Changes in sea levels brought about by Pleistocene glaciations would have joined populations of the numerous adjacent islands on shallow banks such as the Great Bahama Bank during glacial periods and isolated them during interglacial sea level rises, facilitating hybridization and amplifying morphological diversity over multiple eustatic cycles (Harasewych, 2012).
Although the distribution of cerionids throughout most of the islands of the tropical western Atlantic was the result of overland dispersal followed by the break-up of GAARlandia (Greater Antilles + Aves Ridge Land Bridge) and the displacement of Antillean leaks to the Ronheast by the advancing Caribbean Tectonic Plate (Itturalde-Vinent, 2006: fig. 13), the lack of a land con- nection between the islands of the Bahamas and the Greater Antilles precluded overland colonization. Rather, the Bahamian Islands must have been populated by propagules dispersed from the Greater Antilles either by rafting or by hurricanes (Harasewych, 2012: 124). Cle neh (1 938: 495) reported that the fauna of the Bahamas archipelago likely dates only from the Pleistocene and has reached the Bahamas by fortuitous means, noting (Clench, 1938: 484485) that the species of the Great
THE NAUTILUS, Vol. 132, No. 3-4
Page 72 Table 1
Pe ypulation STATION LOCATION LATITUDE
l BP Barker’s Point 24°06.61 N
2» VH Victoria Hill 24°05.83’ N
3 EA E of airport 24°03.85' N
4 Cr N of Cockburm Town 94°03.24 N
5 SU Sugarloaf 24°00.43° N
6 G1234 Sugarloaf 24°00.15° N
7 GB Grotto Bay 23°57.26° N
8 WOQ Watling’s Quarry 23°57.23' N
9 SH1 W of Sandy Hook 23°56.85° N
10 SH2 E of Sandy Hook BST NTT IN
1] rar The Thumb 24°00.S0° N
12 G193b W of Storr’s Lake 24°02.72’ N
13 G194 W of Storrs Lake 24°03.52’ N
14 @1225) Columbus Monument 24°04.06 N
15 G1240 NE of Granny Lake 24°02.89° N
16 CC Crab Cay 24°03.93° N
17 HB Hanna Bay 24°O7.14 N
18 MH Man Head Cay 924°07.54 N
19 NPI North Point 94°07.65° N
20) NP2 North Point 24°07.22’ N
21 GC Green Island (Cay) 24°08.33’ N
Bahama Bank were closely allied to those of Cuba, while the species of the easterly islands (Crooked Island Group to Caicos Island) were affiliated with species from Haiti. He hypothesized that relationships among the terrestrial molluscan faunas of this region are due to island proximity and the paths of hurricanes.
San Salvador is a small island on the eastern margin of the Bahamas platform that has remained isolated by deep water, with little change in island area during the eustatic sea level fluctuations since at least the Pliocene. Its living and fossil Cerion fauna had been spared some of the excesses of early taxonomists, yet has been well studied by modern researchers, particularly in the areas of ecology and chronostratigraphy, due largely to the presence of the Gerace Research Centre (formerly Bahamian Field Station) on the island. The genus Cerion is well represented in the fossil record aft San Salvador, with a documented dated presence spanning the past 140,000 years (Hearty and Schellenberg, 2008).
In the present study, we analyze the morphology of Cerion shells from representative populations spanning the island of San Salvador in order to determine the number of phenotypes and their distribution, and to reconcile these with the type specimens of all taxa de- scribed from San Salvador or Watling Island. A second series of analyses incorporates measurements from a small subset of well-preserved, adult fossil specimens for which ages have been determined (Hearty and Schellenberg, 2008). These analyses follow the chro- nology of patterns of phenotype ‘distribution from the Pleistocene to the Recent to the extent possible given the sampling.
Conclusions of earlier works on the Cerion of San Salvador are reviewed in the context of our findings.
Populations of Recent of Cerion from San Salvador (Watlings Island) included in this study.
LONGITUDE CATALOG NUMBER N GROUP
74°30.87 W USNM 1110076 10 74°31.27 W USNM 1110077 10 74°30.90° W MCZ IP 190243-190252 10 74°32.16 W USNM 590160 10 74°31.89° W USNM 1110078 8 74°32.17 W MCZ IP-190212-190221 10 74°33.63° W USNM 1110079 10 74°32.87 W USNM 1110080 10 74°29.91° W USNM 1110081 10 74°29 32) W USNM 1110082 10 74°27 .35° W USNM 1110088 10 74°27 17 W MCZ IP 190195-190204 10
74°27.19° W 74°25.80° W
MCZ IP 190205-190211 ff MCZ IP 190264-190273 9
[NS) [82) (82) [RS) [N2) [8S) [R2) [8S) [8S) [8S) [SST Ps fe ed) Pt tt (8) 1)
74°27.19 W MCZ IP-190222-190231 10
74°25.80' W USNM 1110087 ) 74°27.03° W USNM 1110085 4 74°26.97 W USNM 1110086 § 74°27.44 W USNM 1110083 8 14°27.37 W USNM 1110084 10 74°30.51° W USNM 359519 10
TAXONOMIC HISTORY
Known as Guanahani to the native Lucayan inhabitants, the island was named San Salvador by Christopher Co- lumbus when he first sighted land in 1492. It was settled by John Watling in the i 7th century and became known as Watlings Island. The name San Salvador was officially transferred from what is now Cat Island to Watlings Island in 1925 as historians concluded that this was the island where Columbus first landed in the New World.
Dall (1894: 117) reported on the molluscan fauna of Watling Island based on material sent to him. He iden- tified as Cerion (Strophiops) glans (Kiister, 1844) a single, rather small specimen taken from about a pint [0.473 L] of beach drift from Watling Island lagoon during a U. S. Fish Commission sampling. Dall (1905: 438) later wrote that no Cerion had previously been noted from Watling Island, and that his earlier report of C. glans from this island was due to an error.
Pilsbry (1902: 265) described the subspecies Cerion eximium fraternum (Figure 1) as being from San Salvador, based on material attributed to Bland. Clench (1938: 531) reported this taxon from Little San Salvador (now also known as Half Moon Cay), an island between Eleuthera and Cat Cay. He noted that the exact type locality for this taxon is unknown, but "as Bland had received other species of Cat Island material from near the center of the island, it is quite probable that his fraternum material came from the same area." As Pilsbry’s publication pre- dated the renaming of Watlings Island, it seems clear that Cerion eximium fraternum was described from Cat Island
(called San Salvador at the time) and does not occur on the island that is known as San Salvador today.
In his report on land snails collected in the Bahamas, Dall (1905: 438-439) proposed two new species, Cerion
=I
ey)
M.G. Harasewych and M.]. Tenorio, 2018 Page
Figures 1-6. eximium fraternum Pilsbry, 1902. Lectotype (designated by Baker, 1963: 206). ANSP ber San Salvador (Bland). 2. Cerion (Strophiops) watlingense Dall, 1905. The specimen illustrated by Dall (1905: pl. 58, fig. 7) is here designated as the eee pe for this taxon. USNM 132970. Watling Island [San Salvador]. 3. Cerion (Strophiops) inconspicuum Dall, 1905. The specimen illustrated by Dall (1905: pl. 58, fig. 2) is here designated as the lectotype for this taxon. USNM 37676. Hs atling Island. 4. Cerion (Strophiops) i inconspicuum lacunorum D: all, 1905. The specimen illustrated by Dall (1905: pl. 58, fig. 4) is here designated as the lectotype for this taxon. USNM 127494, Watling Island, on the shores of the lagoon. 5. Cerion coloni Bartsch, 1924. Lectotype (designated herein), USNM 359377, C ‘olumbus Point (Crab C ay), San Salvador, Bahamas. 6. Cerion rodrigoi Gould, 1997. Holotype, MCZ IP 190264. Windswept terrace of Crab Cay adjacent to the Columbus Monument, San Salvador, Bahamas. Scale bar = | cm for all images.
Ape rtural, late ral, and dors: ul views of primary type spe cimens of Cerion de NYG ribe ole as from San Salv ador. at Cerion
watlingense (Figure 2) and Cerion inconspicuum
specimens during his two weeks on the island. This (Figure 3), both from Watling Island. He also described
published report included a photograph (Bartsch, 1924:
the \ variety Cerion inconspicuum lacunorum (Figure 4) from the shores of the lagoon on Watling Island.
Paul Bartsch conducte d fieldwork on San Salvador in August of 1923, noting in a report of his visit (Bartsch, 1924) that he had gathered approximately 25,000
fig. 41) of seven specimens of Cerion on a leaf and was captioned "Cerion coloni, new species, ... taken at the base of the Columbus Monument, San Salvador." sociates a name with an illustration prior to 1931, it is an available name (ICZN 2012, Article 12.2.7). The taxon
As this as-
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THE NAUTILUS, Vol. 132, No. 3-4
Cerion coloni Bartsch, 1924, (Figure 5) was not recorded in Clench’s (1957) catalog of the Cerionidae, nor in Ruhoff ’s (1973) listing of Bartsch’s zoological taxa. Un- aware of Bartsch’s publication, Gould (1997) described Cerion rodrigoi (Figure 6) from the same locality.
Many eubee quent researchers (e.g., Woodruff, 1978: Poth et al. 2010: Rose, 1990) have regarded the Cerion fauna of San Salvador to consist of a single variable species, while others (e.g., Baldini et al., 2007:174; Hearty and Schellenberg, 2008) have elected to avoid species nomenclature, "as neither proof nor consensus exists with regard to species level taxonomy".
MATERIALS AND METHODS
A geometric Menges analysis of shell shape was performed on 193 specimens of Cerion snails from 21 populations spanning San Salvador Island (Table 1). Figure 7 illustrates the location and a representative specimen from each population.
Specimens were individually photographed in an apertural orientation using a digital camera (Nikon D300) with the lens axis oriented perpe oacltaulle to the coiling axis of the shell. Images were taken at a distance of 35-40 cm in order to minimize image distortion and parallax error. To avoid the effects of Alllennateeay only adult specimens, free of repaired breaks and with a well- -preserved em- bryonic shell and a fully formed terminal adult aperture having an expanded and reflected lip indicative of terminal growth (¢ (Gould and Woodruff, 1978: 381), were selected for analysis.
Shells were digitized using TPSDIG2 (Rohlf, 2005). A total of 24 points were selected to capture shell shape (Figures 8-9). All points were treated as landmarks. These points cover most of the ratios used for classical charac- terization of Cerion shell morphometry (Gould et al., 1974). Taking several photographs of the same speci- men ensured the reproducibility of the measurements by checking that the deviation in landmark positioning was minimal Thus, the same Cerion shell (maximum length 25.56 mm) was photographed 10 times, even with dif- ferent optics in the camera. The maximum error in landmark positioning was 0.13 mm (0.51 % referenced to the size of the shell). The mean error was 0.06 mm (0.23 % relative to the size of the shell). Landmarks were aligned by a standard generalized least squares (GLS) procedure. Procrustes coordinates and centroid sizes (CS) were generated by COORDGENG6bh (Sheets, 2003-2005). CS provides a measure of the geometric size of each speci-
men, and is computed as the square root of the sum of squared distances from each landmark to the centroid of
each specimen’s configuration of landmarks (Zelditch et al., 2004). Partial warp scores (PWS) were derived from the 24 landmarks used for capturing
the shells upon Procrustes alignment and calculation of the consensus reference form. PWS were used as shape variables in our analyses. PWS and their principal components (PC, also known as _ relative warps) were
g the shape of
computed using PCAGENS, which is included in the Integrated Morphometrics Package software suite (IMP 8, Sheets, 2003-2014). CS and the first principal component (PC1) of the partial warps accounted for 92.2% variance in the dataset. Gaussian clustering of these variables (CS and shape PC1 scores) was used to detect population clusters, with the number of clusters determined using the Bayesian Information Criterion (BIC) as implemented i in MCLUST Version 5 software package for R (Scrucca et al., 2016). This program tests different models of clustering, and the fit of each model to the dataset is assessed by means of the Bayesian information criterion.
In our system the best-fit model (BIC —332.56, log L ~132.0738, df = 13, for n = 193) indicated 3 compan or clusters of variable volume and equal shape and ori- entation (VEE model). The pairwise Procrustes distances between groups of populations based upon shell mor- phometry were obtained with TWOGROUPS (Sheets, 2003-2014) using resampling methods (400 bootstraps). The significance of the Goodall’s F-test was assessed by a bootstrappe d F-test. Discriminant Function Analysis (DFA) using PWS+CS as variables, and morphometric grouping as factor was performed with the programs STATGRAPHICS CENTURION XVII and CVAGENS (Sheets, 2003-2014). The latter program was also used for generating the two-dimension (2D) scatter plots and the corresponding deformation grids and vectors. Discrimi- nation was statistically significant along both DF1 and DF2 axes (DF 1: Wilk’s A = 0.0852, y* = 413.8103. df=90, p < 2.22045 x 10°'°: DF 2: Wilk’s A = 0.4458 x = IGS, Gl = WAL ip = DBT x 10").
RESULTS
A geometric morphometric analysis of the shells of 193 specimens from 21 living populations of Cerion from San Salvador Island was conducted. A Principal Component Avnallysi of the PWS yielded two statistically significant components that represent 62.4 % of the overall variation in shape in our sample. Shell height of Cerion from San Salvador is known to exhibit a strongly bimodal distri- bution (Fronabarger et al., 1997). Gaussian clustering of CS and shape PC] data was used to determine the dif- ferent morphometric groups in our sample according to size and shape by the program MCLUST in R. The best-fit model indicated three components or clusters of variable volume and equal shape and orientation (Figure 10). This analysis indicates that the pattern of dtsucbntion of size and shape in our sample of Cerion shells is best represented by three separate groups, partitioned among three different phenotypes. We obtained the initial com- position of each of the groups directly from MCLUST, taking into consideration that the group assignment for specimens falling in the overlap region among clusters may be subjected to increased uncertainty (Figure 11). Final assignment of individuals to each of the three groups was determined by checking the composition of each of the populations against the suggested grouping resulting
M.G. Harasewych and M.]. Tenorio, 2018 Page
ea bank edge and lakes
fe] present shoreline
al +6 m shoreline
LENE
A, Dy. \ i
Figure 7. Map of San Salvador, Bahamas. Locations of sampled populations (solid circles) are identified by alphanumeric station designations (see Table 1). A representative specimen from each station is shown.
from MCLUST Gaussian clustering. Individuals from Group 2, and thus had a higher uncertainty in group each of the populations studied were all members of the classification. same group with the exception of populations VH and CT, Phenotype 1 (Group 1) includes all individuals from
which fell into the region of overlap between Group | and populations: BP, VH, CT, SU, G1234, GB, WQ, SH1, and
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THE NAUTILUS, Vol. 132, No. 3-4
Figures 8-9. coloni Bartsch, 1924, showing 9. positions of the landmarks used in the morphometric analyses.
Position of landmarks. 8. A specimen of Cerion
SH2. Phenotype 2 (Group 2) includes all individuals from populations: TT, G193b, G194, G1225, G1240, CC, HB, MH, NP1, NP2, and GC. Phenotype 3 (Group 3) includes all individuals from a single population, EA.
Phenotype 1 occurs exclusive ly along the western and southern coasts of San Salvador to the arom of the lagoon (Figure 7, BP to SH2). Phenotype 2 occurs along the east coast of the island, from the mouth of the lagoon northward to the base of the North Point Peninsula (Fig. 7, TT to NP2), as well as at several locations inland from the east coast (Fig. 7, G193b, G194, G1240). Populations from an isolated cay (Beane 7 7, C C) off the north coast as well as from the tip of the North Point Peninsula (Figure 7, NP1) include phenotypes primarily from Group 2 but ih several individuals from Group 1, indicating areas of potential overlap or hybridization. Populations
VH and CT appear on the plot clustered in the area of
overlap of between Phenotypes | and 2. The geographic
distribution of these populations in the western coast of the island led to their final classification as members of
Phenotype 1. Phenotype 3 is known only from a single population inland from the west coast (Figure 7, EA).
Confirmation of the composition of these morphom- etric groups was accomplished by means of Discriminant Function Analy sis (DFA). Discrimination was See significant along both DF1 and DF2 axes (Figure 12). A DFA using the above group composition as a factor and PWS+CS as variables classified correctly 190 out of 193 specimens (98.4 %). In the jack-knifed assignment test, the rate was 90.7 % (175 oe out of 193). Using shape- only variables (i.e. leaving CS out of the anelyai) the correct classification rate was 97.4 % (89.1 % in the jack- knifed test). Figures 13 to 24 represent the relativ e po- sition of the specimens from each of the populations in discriminant space.
The three morphometric groups resulting from this analysis differ statistically in size and shape. ANOV A in- dicates that there are significant differences in CS be- tween Group | (mean CS = 35.79) and Group 2 (mean CS = 43.05) (F = 311.37, p = 0.0000), and between Group 2 and Group 3 (mean CS = 34.98) (F = 79.85, p = 0.0000). However, there is no significant difference in size between Group | and Group 3 (F = 080; p = 0!3735): The three groups exhibit significant differences in mean shell shape, as inferred from the calculated pairwise Pro- crustes distances between means and the corresponding results of bootstrapped Goodall’s F-tests (Table 2).
Thus, individuals from groups 1 and 3 do not differ in shell size, but differ greatly in shell shape. The dis- tinguishing features of each of the groups can be extracted fen the shape deformation implied by DF1 and DF2 (Figure 12). Group | is separated from Group 2 along the DF] axis, while € Group 3 is separated from group 1 anal D 2 along the DF 2 axis. Members of Group 2 can be separated from Group | by the larger size of their shells, which are more elongated with a projecting apex and a relatively narrower aperture. Group I is Eiamcrereed by smaller shells, with broadest whorls at mid-length, and a more depressed apex, giving a general oval appearance and with a relatively wider aperture. Members of Group 3 exhibit shell features similar to those of Group 1. However, individuals in Group 3 display a less inflated shell shape, with parallel sides slightly narrowing at the mid-length and with a convex profile towards the apex. She Ils of individuals from Group 1 are relatively broader at the center, with a straight profile towards the more pro- truded apex. The aperture in Group 3 is narrower than in Group 1. Apart from these differences in size and shape, the shells of individuals in groups 1 and 3 also exhibit a more developed sculpture of axial ribs compared to members of group 2. In the latter, the axial ribs may be reduced or even anes nt in certain specimens. This fea- ture provides additional support for the assignment of populations VH and CT to Group 1, given their strong axial ribbing.
Using this morphometric framework, primary type specimens of each of the named taxa from San Salvador (Lectotypes de ssignate od in captions to Figure s 2-5) were digitized and analy zed by means of DFA (Figure 25). The lectotype of Cerion watlingense clearly falls within Group 1 (probability 98. 8%), and is the oldest name available for this phenotype. The lectotype of Cerion coloni and the holotype of C. rodrigoi both fall within
Table 2. Comparison of pairwise Procrustes distances be- tween groups of specimens as defined by shell phenotype, based upon shell morphometry. The significance of the Goodall’s F-test was assessed by a bootstrapped F-test (400 replications).
Group Group Procrustes distance | Goodall’s-F ) I i
] ») 0.0361 30.03 0.0025 1 3 0.0823 31.25 0.0025 2 3 0.0577 16.26 0.0025
M.G. Harasewych and M.]. Tenorio, 2018 Page 77
e =Group1 © =Group 2 e =Group3 wo V9) ros) ei S 3 TS ° i & e Wr SS ae N Nena ee Sune 1 SAate oe @ JPoaS SNe os wis eta y oN ao ac c 0@@ @ 0° o \, C wf pei | fa im Sy \ @ icy 1 ™ @ ° res a) fc) e SQ YH te) re) . _— oo (OS iS : S 2 (WS ! 1 . Sm St CaN 5 ° Na ‘ * }e® 4 1 Nis iF 1F T le a 30 35 40 45 50 CS ° 2 °° N g SSO oo LL nm e ¢ a) oO 7 1,7 = s J a s* s s s 5 s 7 ee eer -3,2 -1,2 DF1
Figures 10-12. Plots of the first principal component (PC1) versus Centroid size (CS). 10. Symbols indicate classification corresponding to the best-fit model as determined by MCLUST. Component means are marked, and ellipses with axes are drawn corresponding to their covariances. 11. Classification uncertainty as determined by MCLUST. Symbol size proportional to degree of uncertainty. 12. Discriminant Function Analysis (DFA) of the shapes and sizes of the shells of 193 specimens corresponding to 21 populations of Cerion snails from San Salvador Island, using Partial warp scores (PWS) + Centroid size (CS) as variables and considering morphometric group as classification factor. The 2D scatter plot of the DF1 and DF2, and the corresponding deformation grids and vectors are shown.
Group 2 (probabilities > 99.9%), with C. coloni being the Group | and Group 3 with the algorithm assigning them oldest available name for this phenotype. The lectotypes to Group 3 at 95 % probability for C. inconspicuum, and of Cerion inconspicuum and Cerion inconspicuum lacu- 51.4 % for C. inconspicuum lacunorum. Cerion incon-
norum have morphologies that are intermediate between spicuum is the oldest available name for this phenotype.
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@ =Groupi 0 9 @ =Group2 ¢ B =Group3 23 o % ° tC) were & ° eo %@ ‘ ry oe MAAS SE, 0.3 $ e N v be ¢ 4 V re e, + % 9 © ee Po ° a ya” Geo oth -17 ° fs. ° a =e vy =P 13 oa @ =TT a a m-EA | -57 5 ~ -52 -~32 -12 0.8 28 48 = @ =Group1 Vv 0 @ =Group2 ¢ @ =Group3 23 * e O49? e oo é AY 1 * e Os 0 03 of 3 ° N 4 : ¢Y v oo ¢ - 8 oe oe +9 e ds 9 } @ o eet ° -17 . fs, ° a 1] -37 15 a v =CT a s @ = 6194 -5.7 5 ~5.2 -3.2 -1.2 08 28 48 sc @ =Group1 7 4? @ =Group2 ¢ m =Group3 23 © %
2¢ 2 © eos e -17 LE A Bo e a of] -~37 \7/ is V =G1234 os r @ =G61240 esi, a] : -52 -32 =-12 08 28 48 43h er Z| = Group 1 7 @ =Group2 ® B =Group3 23 *” % e e we? » ¢ = VASesle e Os 4 © SPowe tye e °% @ 0.3 %, $ oe oy ody a a ¢ We “oe S te Go ° fa) o¢ 4 ) oth ° —f}>/ : e a5 e a C] -37 19 ba Vv =wa 5 a @ =HB -57 5 , -5.2 -32 -12 0.8 28 48
DF1
3 % So @ =Groupi @ =Group2 ¢ m =Group3 23 ms a uke WX Mo” 5 | Gee ¢ 8, e S$ 2c é $,¢ e On Oo 03 0, 3 Ory, ° mM ° th Wr? ¢ PS 00 e ° 2 34 e ote e -17 m4 #3. ° a) -37 al > cts s a =G193b -57 "a z | -52 -32 -12 0.8 2.8 48 ig @ =Groupt one, @ =Group2 ¢ B =Group3 23 "ee & °¢ *e ee ° ¢. oh ° Saee ? ta 4, ¢ e@ e e e 0.3 %, ‘) ° A ? oe ty v7 o%Y, oe eo ds Po ° ¢ (a) a6 a e oot t ° -17 3 ete e "5 -37 16 ba Vv =SU a a @ =G1225 -57 bs] = ~5.2 ~32 -12 08 28 48 ny @ =Group 1 \f \/ @ =Group2 + @ =Group3 23 vo ° ° * “hf ¢ a or) A Oe ¢ © Poe F r) $, ¢ e Oy e 0.3 e, ve o é GO a 0 ons Oey A oy ¢ (a) 24 ¢ g e ogee e -17 $ fs. e a ~37 IB oo 3 =cc ] a @=CCc -57 5 - -52 -32 =i) 08 28 48 *3 @ =Group1 2 © @ =Group2 * @ =Group3 23 w e e o *h® °¢ eo = =%%o i. Oo? Rog Be © S308 & nO of $% % ° °° iN ; . ¢ A é oo? A oS 2 e e DR 4 % e oc8 8 ° -17 % ese e a r] -37 2 ba v =SHi QO. ‘ art =y) 5 -5.2 -32 -12 0.8 28 48 DF1
Figures 13-24. Positions of specimens from individual living (Figures 13-23) and fossil (Figures 22-24) populations in the dis-
criminant space.
A number of well-preserved fossil specimens that had been dated by amino acid racemization in a previous study (Hearty and Schellenberg, 2008) were digitized, scored, and classified by DFA as had been done with the type
material. The oldest well-preserved specimens suitable for morphometric analyses were from French Bay (be- tween WQ and SHI on Fig. 7). All three specimens from SFB4b(1) (Figure 26) (Hearty and Schellenberg, 2008:
M.G. Harasewych and M.]. Tenorio, 2018
a @ =Group 1 0 0 @® =Group2 Vv B =Group3 23 Ww “tw VY 8 Me ° @ 30 ¢ ing Vv eo eas oo “A 03 © g AJ @ ¢ * My ge ue af oe on ¢ We oP GS %s Ze Oo ¢ 4 @ 203% ° 17 A $e. e a a ba v =SH2 a a ® = NPI By -3.2 -1.2 0.8 2.8 4.8 @ =Group1 Sa, @ =Group2 4 ® =Group3 % % @ @ ” a ‘ e : 4g 7 ; oe e Vv ‘ oA 7 % ry 5 nd ee, A N ¢ y $e rr &S 0 C) LL > % ¢ 4 Oo ® 2 $4" Bo og . ® @ 2 + e e a a DB aa Vv =SMHI1b a | @=GC : ay ~ -32 -1.2 0.8 2.8 4.8 DF1
Page 79
a we , & oe © 003% Me ei des TYPE SPECIMENS |
@ =Group1 @ =Group2 @ =Group3 ar ne @ Cane’, 0 Oe. ° ° e V = SFB4b @ =NP2 2.8 48 4.3 @ =Group1 vane, @® =Group2 % B =Group3 2.3 o §% @ e@ we? » °¢ ween e e By \ % iy CBee", 4 03 ) %o N , e ry of, Se Le 4 6% % 9 90 e@ &s Po =) ge” e @ eet e -1.7 V/ & e es © @ Vv @ a a -3.7 D4 g v =STGtc a = ® =SCC4d 57 a -5.2 -3.2 -1.2 0.8 2.8 4.8 DF1 @ =Group1 i @ =Group 2 % B® =Group3
HK = watlingense
*¢ = inconspicuum —|
= /acunorum P #6 = coloni
ef = rodrigoi
0.8 2.8 4.8
DF1
Figure 25. Positions of primary type specimens in the discriminant space.
Fig. 2, Table 2) fell well within Group 1 (Figure 22) (probabilities > 99 %), indicating that this phenotype has been represented on the southern coast of San Salvador between 125,000 and 140,000 years before present. Three
fossil specimens from Man Head Cay SMH 1b (Figure 27) (Hearty and Schellenberg, 2008: Fig. 2, Table 2) also fell well within Group 1 (Figure 23) (probabilities > 99.8 %), indicating that this was the only phenotype occurring on
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Figures 26-29.
Fossil specimens of San Salvador Cerion used in morphometric analyses. 26. Three specimens from the French Bay Member of the Grotto Beach Formation [station
SFB4b(1), Hearty and Schellenberg, 2008: Fig. 2, Table 2],
(125,000— ie 000 years before present). 27. Three specimens non Man Head Cay {station SMH 1b, Hearty and Schellenberg, 2008: Fig. 2, Table 2], (90,000-110,000 years before present). 28. Five specimens from The Gulf [Station STGIcl, He arty and Schellenberg, 2008: Fig. 2, Table 2], (80,000-90,000_ years before pre sent). 29. Thee specimens from Crab Cay [SCC4d (1-2), Hearty and Schellenberg, 2008: Fig. 2, Table 2], (70,000-80,000 years before present).
Man Head Cay between 90,000 and 110,000 years before
present. Modern Cerion on Man Head Cay are all of
Group 2 (Figure 20). Fossil samples STGlel (Figure 28) (Hearty and Schellenberg, 2008: Fig. 2, Table 2 \ ron TN Gulf (very near SH1 on Figure ays were morphologically ai ‘rse (Figure 24), with three falling within Group
1 (probabilities > 99 %), 1 within Group 2 (probability
93 %) and one questionably within Group 3 (probability of
only 61.9 %), indicating that in addition to phenotype 1, phe notypes 2 and possibly 3 were present on San Salvador between 80,000 and 90,000 years before prese! nt, al- though in a region of the island where phenotypes 2 and 3 do not occur Roakey, Three fossil samples from Crab Cay (Figure 29) [SCC4d(1-2), Hearty and Schellenberg, 2008:Fig. 2, Table 2] near the geographic center for
Group 2 in the modern fauna, reveal that both Group 1 (probability > 95 %) and Group 2 (probabilities > 99 %) (Figure 24) were present at this location between 70,000 and 80,000 years before present.
DISCUSSION
Despite minor differences in the number and types of measurements and analytical algorithms, our results concur with prior morphometric analyses (e.g. Hearty et al., 1993; Fronabarger et al., 1997: Hearty and Schellenberger, 2008) in identifying two primary shell phenotypes of Cerion living on the idbarn of San Salvador, one centered along the w estern coast (Group 1), the other along the eastern coast (Group 2) of the island. The presence of a third phenotype (Group 3) with a very restricted inland distribution has not been previously reported, since prior studies had not sampled in the area in which it occurs.
A number of these studies had dealt with morphological changes in Cerion shells throughout the continuous fossil record that spans 140,000 years. Hearty et al. (1993: Fig. 6) had reported that a single phenotype was present on San Salvador in the Grotto Beach Formation (MIS 5e), the Gulf Unit (MIS 5a), and the Man Head Unit (MIS5a), but that a different phenotype was present in the later Almgreen Cay Formation (MIS5a). They noted that the more recent Holocene Rice Bay Formation contained shells with both phenotypes. They had also documented (Hearty et al., 1993: Fig. 7) a significant increase in shell length ‘and especially in shell mah within the Almgreen Cay Formation (70,000—90,000 years before present). Hearty and Sche llenberg (2008) ‘subseque ently reported that change 5 in the morphology of Cerion shells on San
Salvador over the past 140,000 years appear to be con- tinuous and generally directional, and that a trend of increasing gross she ll size characterizes each of the in- terglacial “phase s (MIS 5e, MIS 5a and MIS 1). They also commented that live-collected Cerion have a range in gross morphology that nearly encompasses that of the entire fossil sequence.
In morphometric analyses of living and fossil Cerion shell morphology, Fronabarger et al. (1997) concluded that variation is dependent on ies and suggested that environment affects shell shape. They also noted that
variation seen between living and fossil populations is the same as the variation seen in the geogr aphic distribution of living samples.
These studies interpret the considerable variation in shell morphology, both geographic and chronostrati- graphic, based on the premise that there was but a single colonization of San Salvador by Cerion, and that all var- iation resulted from subsequent differentiation within a single lineage. Rose (1990) attributed the different morphologies to intraspecific variation, with the larger, thicker ribs found on shells that inhabited the canton coast being a response to predation by crabs and rodents. Hearty and Schellenberg (2008) attributed rapid changes
M.G. Harasewych and M.]. Tenorio, 2018
Page 81
in shell size during MIS 5a as responses to environmental factors due to climate change during this interglacial period with its warmer and wetter aiimate: aBuew con- cluded that multiple Cerion extinctions and recoloniza- tions during the past 140,000 years are improbable due to the continuous fossil record, and that multiple species are unlikely to occur on San Salvador.
Gould (1997) noted the intermediate position of San
Salvador he tween the "two major geographic domains of Bahamian Cerion" and concluded that the Cerion fauna of
the island is descended from propagules from both geographic regions. He considered Cerion watlingense (Group 1) to be derived from the "ribby" bank- “edge phenotype (see Gould and Woodruff, 1986) of the Great Bahama Bank, while the "triangular" phenotype, which he named Cerion rodrigoi (a junior synonym of C.
coloni Bartsch, 1924) (Group 3) to have originated in the southeastern Bahamian islands. He speculated that a third phenotype, corresponding to the "mottled", bank interior phenotype of the northern Bahamas, would either be absent or restricted to the interior of San Salvador because the island has no bank-interior coast (a coast that was inland when eustatic sea level fell, exposing larger banks during glacial periods). He commented that it was not known whether the "ribby" and "mottled" phenotypes represent distinct, monophyletic lineages throughout the northern Bahamas, or whether the plienotypes arise in- dependently and convergently as adaptations to common environments.
Our findings of three distinct Cerion phenotypes on San Salvador are congruent with Gould’s hypotheses. Phe- notype | corresponds to Cerion watlingense, derived from a propagule originating on the Great Bahama Bank, has been present on San Salvador since at least 125,000 to 140,000 years before present, and was widespread throughout the island, occurring at Man Head Cay By 90, 000 to 110,000 years before present. Phenotype 2 corresponds to Cerion coloni, and is believed to be de- rived from a propagule originating in the southern Bahamas. Based on our limited sampling of fossil speci- mens, earliest records are known from the southeastern corner of San Salvador between 80,000 and 90,000 years before present, although in a region of the island where this phenotype does not occur today, This phenotype appears to have expanded its range northward along the eastern coastline of San Salvador, here it likely replaced or hybridized with Phenotype 1. It co- occurred with phenotype 1 at Crab Cay between 70,000 and 80,000 years before present, but is the only phenotype present along the east coast in the modern fauna, including at Man lead Cay, which was inhabited exclusively by phenotype 1 between 90,000 to 110,000 years before present. Phenotype 2 continued to expand westward along the northern coast of the island, including to several of the offshore cays. Populations at the northern tip of North Point Peninsula and Green Cay are dominated by Phe- notype 2, although vestiges of Phenotype 1 still remain at these localities. The presence of a third, morpho- logically distinct phenotype (Group 3) on San Salvador,
corresponding to the "mottled" phenotype of the northern Bahamas, is presently known from a single population in the interior of the island, in agreement with Gould’s prediction.
Analyses of shell morphology of living and fossil Cerion occurring on San Salvador may be interpreted in the context of several bioge -ographic scenarios, each with different evolutionary aad systematic consequences. The single introduction hypothesis predicts that all Cerion living on San Salvador are part of a monophyletic clade, and all have a most recent common ancestor not shared with Cerion from other islands. As such, they may all be members of a single species, Cerion watlingense, or the phenotypes may have differentiated sufficie sntly to com- prise either separate species or subspecies of C. watlin-
gense. If the multiple introduction hypothe sis 18 correct,
Cerion watlingense as well as Cerion inconspicuum would be more blosely related to species from the Great Bahama Bank, while Cerion coloni would be closer to species from the southern Bahamas. A molecular phylogeny with ap- propriate sampling should resolve between these two hypotheses and provide insights into the processes by which Cerion achieve and modify their patterns of ex- treme morphological diversity.
ACKNOWLEDGMENTS
We are grateful to Dr. Paul Hearty for his extensive ltdkaordk on San Salvador [supported by NSF award EAR 0106936 (Goodfriend, Gould, and Harasewych PI’s) through a subcontract to Dr. Hearty], and for depositing the wacker material for his published study, including dated fossil specimens, in the collections of the Netiomell Museum of Natural History, Smithsonian Institution. We also thank Jessica Cundiff and Adam Baldinger of the Museum of Comparative Zoology for alan available specimens in their care. We chen the reviewers, Dr. James Carew and Dr. Timothy Pearce, for their helpful comments and suggestions.
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THE NAUTILUS 132(3-4):83-90, 2018
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How many micromollusks are there? A case study on species richness in Hawai‘i, with the description of a new species of Murdochella (G astropoda: Epitoniidae)
Bianca Campagnari
Quasars to Sea Stars Teen Program
Santa Barbara Museum of Natural History
2559 Puesta del Sol, Santa Barbara CA 93105 USA biancacampagnari@gmail.com
Daniel L. Geiger
Department of Invertebrate Zoology
Santa Barbara Museum of Natural History
2559 Puesta del Sol, Santa Barbara CA 93105 USA dgeiger@sbnature2.org
ABSTRACT
Four statistical metrics (Chaol, Chao2, ACE, and rarefaction) were used to estimate the total number of species of shelled micromollusks based on composition of nine benthic samples collected from the sublittoral zone off Wailea, Maui, Hawai'i. There was a total of 250 species in the nine samples analyzed, and the estimated total number of species of shelled micro- mollusks in this area, based on the. statistical metrics, was 317-375 species. Murdochella eee new species is de- scribed and Pelycidion habei (Kay, 1979) is discussed.
Additional Keywords: Biodiversity, endemism, oceanic island, Indo-West Pacific
INTRODUCTION
This study estimates the species richness of shelled marine micromollusks off of Wailea, Hawai'i. Micro- mollusks are defined as having shells smaller than 5 mm as adults (Geiger et al., 2007). There are only 1,300 named species of shelled marine mollusks in Hawai'i (Severns, 2011), out of an estimated 37,000 species in the entire Indo-West Pacific (Contrafatto and Minelli, 2011). This difference is likely because of Hawaii's geographic isolation. Biodiversity in the Indo-West Pacific peaks around the islands of Southeast Asia, and decreases outward from there (Severns, 2011). Endemism of Hawaii's malacofauna is 21%, which is the world’s highest. Severns has recorded 394 micromollusk species (Severns, 2011). Estimates for the total number of named mollusk species in the world reach 70,000-76,000, while estimates including undescribed species go as high as 200,000 (Rosenberg, 2014).
The shells in this study were part of a thanatocoenosis, or death assemblage; the composition of the samples is assumed to represent the composition of the surrounding areas. We used non-parametric calculations of species richness, which estimated the total number of species
based on the relative abundance of rare and common species in the sample. An area from which a sample with a relatively high number of rare species was taken is esti- mated to ene ve a high number of unfound species, because a high number a rare species indicates high species heterogeneity and low sampling intensity (Gotelli et al., 2007).
MATERIALS AND METHODS
Nine dredged samples from six locations off Wailea, Maui, Hawaii, at depths of 85-140 meters were collected and kindly made available by M. Severns (Figure 1, Table 1). The bulk sediment samples were sorted, and all speci- mens were deposited at the Santa Barbara Museum of Natural History (SBMNH: Table 1); some paratypes were also deposited in the Bernice P. Bishop Museum, Hon- olulu, Hawai'i, USA (BPBM). Identifications were based primarily on Kay (1979), Okutani (2000), Severns (2011), and Okutani (2017).
A rarefaction curve was generated to display the av- erage species accumulation of each sample and was ex- trapolated to an estimate for the species richness of the entire area using the iNEXT function from the R package (Chao et al., 2016; Hsieh et al., 2016). Species richness estimators used in this study were ACE (abundance-based coverage estimate) (Chazdon, 1998), as well as Chao] and Chao2 (Chao, 1984). ACE and Chaol combine all the samples, while Chao2 considers each sample in the study separ ately and constructs an estimate by comparing the samples. ‘In the R package Vegan (Oksanen et al., 2017) the estimateR and spe cpool functions estimated C heal, ACE, Chao2 and their standard errors, respectively.
Images of select shells were taken by scanning e Jectron microscopy (SEM). Standard met thods were applied for uncoated specimens (Geiger et al., 2007). Light micro- scope z-stack images were taken using a Zeiss Axioskop 2plus compound microscope (Zeiss, Oberkochern, Germany). A Cognisys (Traverse City, Michigan, USA)
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THE NAUTILUS, Vol. 132, No. 3-4
Figure 1
ste pping motor connected to a C ognisys StackShot con- troller and programmed through ZereneStacker (Rich-
land, Washington, USA) was couple d to the fine focus of
the microscope with a custom adapter (Aben Machine Products, Canoga Park, California, USA). Step size was calculated based on depth of field of numerical aperture (2.4-152 um). Images were acquired with a Canon 5DsR digital SLR camera; RAW files were processed with DxO Optics Pro (DxO Labs, Paris, France), and stacked with ZereneStacker using the Pmax algorithm. Further digital imaging was carried out in AffinityPhoto (Serif Ltd., Nottingham, Uk).
RESULTS
Diversity Estimates: A total of 4,233 shells belonging to 250 discrete morphospecies were recovered from the samples.
The estimates that used abundance data for the total number
of species in the area encompassed a 95% confidence interval of 288-404 species (rarefaction, ACE, Chaol: Figure 2, Table 2). All three abundance estimates had overlapping 95% confidence intervals, which suggests that the different methods of estimating species richness were consistent, and
Map of Maui. Markers indicate locations of samples.
thus more likely to be accurate. On the order of three- quarters of species of micromollusks in the overall sampled area were represented in the individual samples.
To explore the possible correlation between number of specimens and species in the sample and diversity esti- mates, _ E and C haol were applie .d to each of the nine samples (Table 3). Estimated species richness correlated positively with aan size, but that correlation was in- significant C < 088. p >> 0.1: Figure 3). Similarly, de »pth was found to have no effect on number of speci- mens found (r = 0.06, p >> 0.1: Figure 4).
New Records: The Hawaiian malacofauna has recently been treated by Severns (2011), including 394 micro- mollusks. Although Hawaii is among the best studied tropical areas aril despite it being a rather low diversity region relative to its tropical keine, the faunal inventory is far from complete. Exciting finds such as the highly distinctive Severnsia strombe vlna Geiger, 2016 highlight the potential for new species discoveries, particularh ly in micromollusks (Geiger, 2016). Some additions to the Hawaiian malacofauna stemming from the present study are detailed below.
B. Campagnari and D.L. 2018
Geiger,
Page 85
Table 1 waypoints. Coordinates
Sample site Location
WP267-424 2.5 mi W off Wailea 20.673° N, 156.482° W WP502-506 3.6 mi W off Wailea 2(0).689° N, 156.500° W WP714-716 1.5 mi NE of Molokini 20.651° N 156.479° W
WP384-394 1.5 mi W of Wailea 20.670° N 156.467° W WP267-468 2.3 mi W of Wailea 20.673° N 156.482° W WP267-496 2.3 mi W of Wailea 20.673° N 156.482° W
20.660° N 156.494° W 20.660° N 156.494° W 20.673° N 156.482° W
mi W of Wailea mi W of Wailea mi W of Wailea
WP404-640 WP404-713 WP641-642
CO OO UW OV Ot Ot
SYSTEMATICS Epitoniidae Berry, 1910 (1812)
Murdochella Finlay, 1926
Diagnosis: Shell high-spired, many fine, low, axial la- mellae, crossed by few spiral cords. Base flattened. Pro- toconch bulbous, smooth.
Type Species: Scala levifoliata Murdoch and Suter, 1906, by original designation.
Remarks: The genus is currently known from New Zealand (M. alacer Finlay, 1926, M. levifoliata) Australia (M. macrina Iredale, 1936), Antarctica (M. antarctica Dell, 1990), South Africa (M. crispata Kilburn, 1985, M. lobata Kilburn, 1985), and Alaska (M. turritelliformis Brown, 2018).
400 400 = 300 300 bent Sg 5 % 200 200 3 Q. DN
—_ = — oS
100
0 0 5000 10000 15000 20000 25000 300000 Number of Individuals Figure 2. Interpolated (solid line) and extrapolated (dashed
line) rarefaction curve of all samples. The shaded region is 95% confidence interval. The asymptote is at 350 species, 95% confidence interval covers 305-394 species.
Depth SBMNH
101 m 424489424607, 424609, 453258
$588 m 265846—-265857, 265859—265934
120-125 m 266363-266380, 452777-452856
92m 26625 1-266296
88 m 266081—266181
88-92 m 266008, 265984—266007, 266009—266046, 966048—266054
139-141 m 266297—266333
139-140 m
266334—266362, 424916, 424917,
453262
Sample sites of this study. SBMNH: range of SBMNH registration numbers for the particular site. WP: Severns GPS
124-132 m
454728454731, 454733454738, 454740—454750, 454752454762, 454764, 454765, 454767—4548 15, 454817454819, 454859454915, 457572
Murdochella tricingulata new species (Figures 5-14)
Description: Shell high-spired, holotype 2.7 mm (Figures 5-6), to 4.5 mm (Figure 9). Shell off-white to dark-tan. Protoconch bulbous, smooth. Up to ten tele- oconch whorls. First whorl with more widely spaced, stronger axial cords, no spiral sculpture (Figures 6, 8). Second whorl with spacing and strength of axial lamellae as on remainder or teloconch, peripheral spiral cord distinct, adsutural spiral cord indistinct (Figures 5, 7, 9-11, 13, 14). Subsequent whorls with three spiral cords: strongest to just below periphery, second at two- thirds towards suture, last just above suture. Height of lamellae variable, some specimens with irregular distal horizontal extensions towards anterior (Figure 12). Base concave with pericolumellar cord. Aperture subquadratic.
Type Material: Holotype SBMNH 266361; Paratypes: SBMNH 266300 (two): Severns WP 404-640 (one) (all from type locality); SBMNH 452826 (nine); BPBM 284613 (two): Severns WP 714-716 (three), all from 2.78 km NE Molokini, Maui, Hawai‘i, USA, 20°39.047' N, 156°28.759' W, 131-136 m; SBMNH 454728 (five): Severns WP 641-642 (two), all from off Wailea, Maui, Hawai'i, USA, 20°39.29' N, 156°29.56’ W, 136-144 m.
USA,
Type Locality: Off Wailea, Maui, Hawai‘,
20°29'38" N, 156°2938" W, 152-154 m.
Etymology: Latin tri- = three, -cingulata referring to the spiral cords; noting the three spiral cords.
Remarks: Murdochella tricingulata is most similar to M. levifoliata with the very tight axial lamellae and strong, stepped spiral cords in the lower half of the whorl. The Hawaiian species is overall more slender, and has three distinct spiral cords, as compared to two in M. levifoliata. Both M. antarctica and M. macrina have more rounded whorls with less prominent spiral cords (Dell, 1990;
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Table 2.
Species richness estimates (95% confidence intervals) based on rarefaction curve, ACE, Chaol, and Chao2.
All specimens combined
Separate samples
Species in samples Rarefaction curve
250 350 (305-394 ) 343
ACE Chaol Chao2 (323-363) 346 (288-404)
425 (341-509)
Beechey, 2008; Engl, 2012). The South African (Transkei) M. crispata has Sea nette sculpture with spiral cords well above the periphery of the whorls, while M. lobata has fewer but strong axial lamellae but no spiral cords (Kilburn, 1985). Murchochella turritelliformis Brown, 2018 from the Aleutian Islands, Alaska (Brown, 2018), has a proportionally wider shell and more numerous spiral cords.
The new species exhibits a moderate degree of vari- ability in the sculpture, which appears pradual/clinal among the 19 specimens available. Acoondting, no fur- ther taxonomic separation is justified. The lamellae can be rather low (height less than half distance between la- mellae) to rather high (height greater than distance be- tween lamellae). The specimens with higher lamellae have trapped debris between the lamellae, which cannot be removed by sonication. The specimens with high lamellae tend to deve ‘lop extremely fine and irregular etl hori- zontal extensions towards the anterior part of the shell. Those extensions are always pure white in color, con- trasting with the off-white to dark-tan background color. The specimens with higher lamellae also appear slightly
broader. As this difference in shape can be accounted for by the projecting sculpture, the shape of the main body of
the shell is ve ry similar in all specimens.
Pelycidion habei (Kay, 1979) (Figures 15-19)
Remarks: Severns (2011: pl. 36, figs 8-9) showed two putative species of Pelycidion from Hawai'i i, the holotype of P. habei and Pelycidion sp. They differ nominally in the glassy (P. habei) vs. opaque white (P. sp.) shell color, the degree the whorls are inflated or incised at suture, re- spectively, and the apparent smooth (P. habei) vs. finely clathrate (P. sp.) shell sculpture. The illustrations in Kay (1979: 27, i-j) reveal fine clathrate sculpture, which
Table 3.
suggests that the apparent differences in the images in Severns (2011) are either due to specimen condition or photographic technique, but that they are not systemat- ically relevant. The three specimens illustrated by SEM here ( Figures 15-17) show quite a bit of variability with respect to degree of incision of the whorls are in- termediate between the specimens figured by Severns (2011). It suggests that there is only a single species of Pelycidion in Hawaii.
DISCUSSION OF DIVERSITY
All diversity measures are estimates, and one needs to be aware of their inherent limitations. While an estimator based on incidence data such as Chao2 can be useful because it takes distribution heterogeneity into account, in this study estimators using lnmadenee data such as ACE and Chaol were more use ‘tal because of the small number of samples analyzed and the fact that the samples in this study were of different sizes. The Chao2 estimate is known to stabilize above 30 samples (Gotelli, 2011), while this study was based on only nine samples. Regardless, even though the Chao2 estimate is much larger Siva the others, the Chao2 confidence interval includes the other estimates.
All models used in this study are minimum richness estimators (O’Hara, 2005). Richness estimates of a par- ticular area have been found to be correlated with sample size, suggesting underestimation due to low sample size Cmitroras et ail, 201 3). Here, this correlation was positive but insignificant (r 4 & (N.S. 1) >> 0.1, Figure 3). Because of the correlation between richness ential and sample size, estimates are ideally compared between samples of equal numbers of individuals from different locations to compare species richness (Chao, 2016). The sample s in this study had unequal numbers of specimens (66-650), though the raw sediment samples had approximately the
ACE and Chaol species richness estimates and standard error for each sample. WP: Severns GPS waypoints.
Number of specimens Number of species ACE ACE 95% Chaol Chaol 95%
Sample Site in sample in sample estimate confidence interval estimate confidence interval WP267-624 196 UU 85 75-95 fol 73-89 WP502-506 102 56 123 109-137 110 54-166 WP714-716 432 64 87 77-97 90, 60-120 WP384-394 195 37 i, 62-82 61 33-89 WP267-468 617 $4 129 117-141 131 87-175 WP404-640 66 28 40) 36-46 39 23-55 WP404-713 1 30 51 47-55 43 25-61 WP64 1-642 650 96 115 105-125 12] 95-47 WP267-496 294 57 126 112-140 163 49-277
B. Campagnari and D.L. Geiger, 2015
Page 87
180
160
120
Richness
40 a
20
0 100 200 300 Specimens
Figure 3.
400 500 600 700
Species richness estimates of samples over number of specimens in sample. Statistically insignificant positive correlation
(r < 0.38, p >> 0.1). Diamonds: ACE estimate. Squares: Chaol estimate. Solid line: Chaol regression. Dashed line: ACE regression.
same volume and were collected within kilometers of each other. Depth (88-141 m) as a factor explaining those differences could be ruled out (1? OS, jo» SS Ol, Figure 4). The observed differences appear to reflect true environmental heterogeneity.
The estimated number of species for each sampling site was 62-88% lower than that for the whole area, which shows high distribution heterogeneity. The sum of sample species aighmness is much greater than the total species richness due to many common species among samples. Estimating species richness is known to be more difficult for highly “ay erse locations (Chao, 2016), such as Maui. Comparison of the data presented here is challenging, because of the absence of comparable studies at other localities. An environmental impact report (Russo et al.,
2018) from a waste water treatment outfall site at 33 m off
Oahu listed 165 species of mollusks of all sizes from
700 600 S00 400
300
Specimens
200 @ ¢
100 °
80 390 100 110
Figure 4.
Linear regression of mean depth vs number of specimens found (
approximately 4,500 individuals. Samples were collected by SCUBA diver and small cores, hence, included the full size-range of organisms. Diversity estimators calculated from Ihe raw ate were highly consistent (this study). Chaol estimates a species idhness of 193+12SE, 95% confidence interval 169-217, and ACE estimated 197+ TSE, 95% confidence interval 183-211 (this study). The different size compositions of the two studies make direct comparison difficult. Based on the species list, we esti- mate that less than half the species in Russo et al. (2018) are of comparable small size as in our samples from Maui; that proportion is conservative and in rough agreement with other studies (Aravind et al., 2008; Middlefart et al., 2016: see also below). Accordingly, the contrast of of species richness of micromollusks alone is even more remarkable: Oahu ~80 found, ~100 estimated; Maui 250 found, ~350 estimated. These differences suggest
120 130 140 150
Depth (m)
r = 0.06, p >> 0.1).
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THE NAUTILUS, Vol. 132, No. 3-4
Figures 5-14. teloconch. 7-12. Paratypes SBMNH 45472
Murdochella dicate s new Me cies. 5-6. Holotype SBMNH 266361. 5. Shell. 6. Detail of protoconch and early
7. Shell of small s specimen with low sculpture. 8. Protoconch of 7. 9. Largest specimen
known (4.2 min). 10. SEM image. 11. oe mab of 10. 12. Enlargement of lamellae with irregular distal horizontal extensions towards anterior. 13-14. Paratype SBMNH 452826. 13. Dark-colored specimen with strong axial lamellae. 14. Lighter colored specimen with
lower axial lamellae. Scale bars shells = 1 mm. Scale bars details
environmental heterogeneity and that the sites sampled in this study are more tage rse than the site sampled in Oahu. Differences in de »pth may account for the differ- ence in diversity, as the Oahu samples were taken at 33 m and the Maui samples were taken from 85-141 m. An- thropogenic effects were discounted by Russo et al. (e018) as similar species composition and numbers of species were found in control sites.
Bouchet (2002) recorded 864 species smaller than 4.1 mm off Koumac, New
Caledonia. The higher
100 pm.
number of species found in New Caledonia is probably due to the fact that the sampling area was larger (295 km? vs 18 km?) with greater habitat diversity, and the fact that New Cale Henin is less isolated than Maui. The absolute latitudes of both areas are similar (Koumac 20.5°S, Maui: 20.7°N).
Many micromollusks are missed in habitat assessments because they are smaller than the holes in the mesh used to sort shells out of a sand sample. Additionally, some micromollusks require the use of a scanning electron
B. Campagnari and D.L. Geiger, 2015
ace BC Page 89
Figures 15-19. Pelycidion habei (Kay, 1979). SBMNH 454742. 15-17. Shells. 18. Protoconch of 15. 19. Enlargement of body
sculpture from 15. Scale bar shells = 1 mm. Scale bars details =
microscope to observe distinguishing characteristics (Middlefart et al., 2016). Despite relatively few studies on micromollusks, they appear to be very common. Of the 269 recorded gastropod species of the Western Ghats,
40% were micromollusks (Aravind et al. 2008), and 33% of
recovered species of mollusks from a study in New Caledonia were micromollusks (Middlefart et al., 2016).
The present study shows that even a rather well-studied area such as Hawai'i still harbors a significant percentage of undescribed species. Given Blawentt s high rate of ma- rine endemism, it highlights the need for strong con- servation measures to protect the yet to be recognized biodiversity.
ACKNOWLEDGMENTS
BC was supported in part by a generous donation from Laura Francis, Education and Omirendh coordinator at NOAA Channel Islands National Marine Sanctuary, and the Sea Forward Fund. Jenna Rolle (SBMNH) kindly provided insights into diversity estimates. Marta deMaintenon and Cynthia Hunter helped with constructive comments to improve the manuscript and provided reports on samples from Oahu. José H. Leal provided editorial guidance. Regina Kawamoto (BPBM) provided the registration number for one paratype lot.
LITERATURE CITED
Aravind, N.A., R.K. Patil, and N.A. Madhyastha. 2008. Micro- molluscs of the Western Ghats, India: diversity, distribution and threats. Zoosymposia 1: 281-294.
Beechey, D. 2008. Murdochella macrina. http://seashellsofnsw.org. awEpitoniidae/Pages/Murdoche Ha_macrina.htm [accessed 8/ 28/2018]
Bouchet, P., P. Lozouet, P. Maestrati, and V. Héros. 2002. Assessing the magnitude of species richness in tropical
100 um.
marine environments: exceptionally high numbers of molluscs at a New Caledonia site. Biological Journal of the Linnean Society 75: 421—436.
Brown, L.G. 2018. New species of Nystiellidae and Epitoniidae (Mollusca: Gastropoda) from the northeastem Pacific, Mol- luscan Research. DOI: 10.1080/13235818.2018.1433956
Chao, A., R.K. Colwell, C. Lin, and N.J. Gotelli. 2009. Sufficient sampling for asymptotic minimum species richness esti- mators. Ecology Society of America 90: 1123-1133.
Chao, A., T.C. Hsieh, and K.H. Ma. 2016. iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill num- bers). Methods in Ecology and Evolution 7: 1451-1456.
Chao, A. 1984. Nonparametric estimation of the number of classes in a population. Scandinavian Journal of Statistics 11: 265-270.
Chazdon, R.L., R.K. Colwell, J.S. Denslow, and M.R. Guariguata. 1998. Gitisticd methods for estimating species aries of woody regeneration in primary and secondary rainforests of aortieostermn Costa Rica. In: Forest Biodiversity Research, Monitoring and Modeling. F. Dallmeier and J.A. Comiskey (eds.). Parthenon Publishing, W ashington D.C., pp. 285-309.
Contrafatto, G. and A. Minelli 2011. Biological Science Fun- damentals and Systematics. EOLSS Publishers, Oxford, 331 pp.
Dell, R.K. 1990. Antarctic Mollusca. The Royal Society of New Zealand Bulletin 27: 1-311.
Engl, W. 2012. Shells of Antarctica. ConchBooks, Hackenheim. 402 pp.
Geiger, D.L. 2016. Severnsia strombeulima n. gen. & sp. from Hawai'i (Mollusca: Gastropoda: Caenogastrpoda: Eulimi- dae). Zootaxa 4084: 587-589.
Geiger, D.L., B.A. Marshall, W.F. Ponder, T. Sasaki, and A. Warén. 2007. Techniques for collecting, handling, preparing, storing and examining small molluscan speci- mens. Molluscan Resear 97: 1-50.
Gotelli, N.J. and R.K. Colwell. 2011. Estimating species richness. In: Biological Diversity. A.E. Magurran and B.J. McGill (eds.). Oxford Biology, Oxford, pp. 39-54.
Hsieh, T.C., K.H. Maand A, Chao. 2016. iNEXT: iNterpolation and EXT rapolation for species diversity. R package version 2.0.12.
Kay, A. 1979. Hawaiian Marine She lls. Bishop Museum Press, Honolulu. 653 pp.
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Middlefart, P.U., L.A. Kirkendale and N.G. Wilson. 2016. Australian tropical marine micromolluses: an overwhelming bias. Molecular Diversity Preservation International 8: 17.
O'Hara, R.B. 2005. Species richness estimators: how many species can dance on the head of a pin? Journal of Animal Ecology 74: 375-386.
Oksanen, J., F.G. Blanchet, M. Friendly, R. Kindt, P. Legendre, D. McGlinn, P.R. Minchin, R. B. O'Hara, G.L. Simpson, 2. Solymos, M. Henry, H. Stevens, E. Szoecs and H. W agner. 2017. vegan: Community Ecology Package. R package version 2.4-4. https://C RAN.R- -project. org/package =vegan
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THE NAUTILUS 132(3-4):91—100, 2018
ace Page 91
Catalog of the taxonomic updates of northeastern Pacific Late Cretaceous shallow-marine bivalves and gastropods named from
1874 to 1966
Richard L. Squires
Department of Geological Sciences
California State University
Northridge, CA 91330-8266, USA
and
Invertebrate Pale ontology!
Natural History Museum of Los Angeles C Jounty Los Angeles, CA 90007 USA
richard.sq uires@csun.edu
ABSTRACT
Published taxonomic updates of northeastern Pacific (southem Alaska to Baja Califomia, Mexico) shallow-marine Late Creta- ceous bivalves and gastropods named between 1874 and 1960, are catalogued for the first time. Of the 143 species (including “varieties,”) 68 are bivalves and 75 are gastropods. These species were named prior to the usage of more effective paleontologic procedures initiated by Willis P. Popenoe in 1957. The junior homonym Inoceramus pacificus Anderson and Hanna, 1935 is renamed as Inoceramus georgeedavisi new name Squires.
Additional Keywords: Vancouver Island, California, Inoceramus
INTRODUCTION
The purpose of this paper is to catalog, for the first time, the taxonomic name changes of shallow-marine species of Late Cretaceous bivalves and gastropods, which were named by early molluscan paleontologists (between 1874 and 1966) in the region extending from southern Alaska to northern Baja Calligoveatn Mexico. This region corresponds to the “Northeast Pacific Subprovince (NEP)” of Kauffman (1973: fig. 1). Squires (2018) provided a discussion of the geologic setting and paleoclimate conditions of this paleo-biotic subprovince. Stemming from his field work in California in the 1860s, William M. Cabb was the first paleontologist to collect and name Cretaceous shallow-marine mollusks from the NEP region. His monumental contributions were catalogued hee Groves and Squires (2018). Subsequent NEP pale- ontologists during the late 1880s to the late 1950s were, in alphabetical order, Frank M. Anderson, Ralph Arnold, James G. Cooper, Donald H. Dailey, E. B. Hall and Arthur Ambrose, Roy D. McLellan, Charles W.
] . Research Associate
Merriam, Michael A. Murphy, Earl L. Packard, Willis P. Popenoe, Peter U. Rodda, Clarence A. Waring, Charles A. White, and Joseph F. Whiteaves. Although Anderson’s Late Cretaceous monograph was published in 1958, his research was actually done in the 1940s. He died in 1945, and the volume was published posthumously 13 years after his death.
Starting in the late 1950s, there was a fundamental transformation in how NEP Late Cretaceous bivalves and gastropods were studied. Popenoe (1957: 430) imple- mented a method that relied mainly on his own large and well-curated collection of fossils: a collection now re ssicling at the Natural History Museum of Los Angeles County Museum of Natural History. “His method of careful collecting, the recording of precise geographic locality and stratigraphic position, ‘and accurate identifications has resulted in so much more orderly knowledge of the Late Cretaceous molluscan faunas that it now seems impossible that these faunas could have been in such confusing disarray as when he began his studies” (Saul et al., 1989: 117). Popenoe selected a representative number of speci- mens from each major formation in which they occur and then carefully cleaned the hinges of bivalves and the ap- ertures of gastropods. C Careful “deamon of these critically important morphologic areas allowed fora much better basis for the systematic placement of taxa. He used detailed biostratigraphy to more accurately show lineages of species and their evolutionary trends. Initially, he waltts d only on ammonite data for biostratigraphic control but later (Popenoe et al., 1960) added inoceramid-bivalve zones. Saul, who was mentored by Popenoe, perfected the method begun by him and gr eatly helped to introduce it to current NEP molluscan paleontologists. The method relies on new biologic and stratigraphic data in order to keep everything updated. In the last 20 years or so, there have been. many updated systematic and biostratigraphic studies of NEP
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bivalves and gastropods, and these studies are the basis of this present paper. It is relevant to mention that species named after the onset of Popenoe’s transformation are much less prone to being changed, than those named prior to it. There are, nevertheless, many other bivalves and gastropods, named prior to 1960, which are in need re-investigation. More name updates will be made in the future.
MATERIALS AND METHODS
The information was largely extracted from the literature. Study area papers concerning Late Cretaceous bivalves and gastropods but without any taxonomic updates were not included. Cold-seep taxa are not included. The au- thors are listed in alphabetical order. Bivalves are listed before gastropods, but a few of the listed species concern only gastropods. The species names, which are also in alphabetical order, include the parenthesized genus name in which the author originally placed the species, along with its date, page, figure number(s), and holotype number, unless otherwise noted. The museum. type- specimens numbers were mostly gleaned from the cited literature. The CASG numbers were derived from its in-house records. The USNM numbers were derived from the USNM online database [natural history.si.edu/ rc]. All of the “original” information is followed by the updated name (in bold) and the currently known ge ologic age, geographic range (listed from north to south), and FER oReOS (s) concerning the updates of the species. For
the geologic age, standard (European) geologic stages of
the Late Cretaceous are used, with the sequence from older to younger being Cenomanian, Turonian, Con- iacian, Santonian, Campanian, and Maastrichtian. Abbreviations used in conjunction with type-specimen numbers are: CASG: California Academy of Sciences, San Francisco; GSC: Ge ological Survey of Canada, Ottawa, Ontario; LACMIP: Neveu History Museum of Los Angeles County, Invertebrate Pale ontology Department, Los Angeles: LSJU, Leland Stanford Junior University, Sramiiorl California [collection now at CASG]; UCMP: University of California, Berkeley, Museum of Paleon- tology,; UO, University of Oregon, Eugene, Oregon; USNM, National Museum of Natural History, Smithso- nian Institution. Abbreviations used for geographic areas are: BX (Baja California, Mexico), CA (California), QC (Queen Charlotte Islands, British Columbia, VI (Van- couver Island area, British Columbia; including adjacent islands along eastern coast), WA (Sucia Island, Washington).
ANDERSON
Bivalves
anaana (Pholadomya) Anderson, 1902: 73, pl. 7, fig. 151; plastohypotypes UCMP 30251, 30252. Liopistha anaana (Anderson, 1902), Turonian, CA. Popenoe (1937: 384), Anderson (1958: 119).
banosensis (Glyc Yes) Anderson, 1958: 98-99, pl. 73, figs. 1-3; CASG 28310.01. Glycymerita banosensis
(Anderson, 1958), late Campanian to latest Maas- trichtian, CA, BX. Squires (2010: 908).
bowersiana (Trigonia) Anderson, 1958: 116, pl. 26, fig. 8 CASG 252.01. Musagonia bowersiana (Anderson, 1958), Turonian, CA. Cooper (2015: 27); no age update found.
branneri (Trigonia) Anderson, 1958: 112, pl. 17, fig. 5; UO 26859. Louella fitchi (Packard, 1921). Cenomanian/ Turonian boundary, OR, CA. Saul (1978: 53), Cooper and Leanza (2017: 330).
buttensis (“Trigonocallista”) Anderson, 1958: 140, pl. 59, figs. 1, la; CASG 27838.15. Calva (Egelicalva) but- tensis (Anderson, 1958), early Campanian, AK, VI, CA. Saul and Popenoe (1992: 36).
churchi (Trigonia) Anderson, 1958: 115, pl. 17, figs. 6, 7 CASG 1788.01. Notoscabrotrigona evansana (Meek, 1858), early to late Campanian, VI, CA. Jones (1960: 436), Cooper (: Ke BS).
colusaensis (Trigonia) Anderson, 1958: 110, pl. 1, fig. 6; lectotype UCMP 12171. Yaadia leana Gabb, 1876, early Turonian to Coniacian, OR, CA. Saul (1978: 33).
contracostae (Inoceramus) Anderson (1958: 103, pl. 18, figs. 3, 4), CASG 29084.01, two forms: a a Coniacian species of Sphenoceramus? and a Coniacian? to Santonian species of Mytiloides?, CA. Kauffman (1977: 189).
duplicostatus (Inoceramus) Anderson, 1958: 100, pl. 17, figs. 3, 4, CASG 61724.02. Mytiloides duplicostatus (Anderson, 1958), Turonian, CA. Kauffman (1977: 189).
eolobatus (Inoceramus) Anderson, 1958: 99, pl. 18, fig. 13; CASG 1291.01. Inoceramus crippsi? Mantell, 1822, early to mid Cenomanian, CA. Kauffman (1977: 188).
gabbiana (Mactra) Anderson, 1902: 74, pl. 7, fig. 156; CASG 1. Cymbophora gabbiana (Anderson, 1902), Turonian, CA. Saul (1974: 1084).
glennensis (Inoceramus) Anderson, 1958: 99, pl. 17, figs. 1, 2; CASG 61629.01 Mytiloides sp., most likely M. opalensis elongata (Seitz, 1934), early Turonian, CA. (Kauffman, 1977: 188).
hemphilli (Trigonia) Anderson, 1958: 115, pl. 52, figs. 9, Ya, 9b. CASG 994. Popenoella hemphilli (Anderson, 1958), early Maastrichtian, CA. Saul (1978: 50), Cooper and Leanza (2017: 329).
jacksonensis (Inoceramus) Anderson, 1958: 100, pl. 43,
figs. 1, 2; not in CASG database. Inoceramus crippsi? Mantell, 1822, early to mid Cenomanian, OR. Kauffman (1977: 188).
klamathensis (Inoceramus) Anderson, 1958: 104, pl. 18, figs. 1, 2; syntypes CASG 61914.01, 61914.02. Sphe- noceramus lingua? (Goldfuss, 1836), Coniacian? to Campanian, CA. Kauffman (1977: 186).
klamathonia (Trigonia) Anderson, 1958: 112, pl. 30, fig. 4., CASG 61873. Notoscabrotrigona klamathonia (Anderson, 1958), Turonian. Cooper (2015: 25), CA.
meekianus (Inoceramus) Anderson, 1958: 101, pl. DO). figs. 5, 6; CASG 228.01. Mytiloides meekianus (Anderson, 1958), Coniacian?, WA. Kauffman (1977: 180).
pentzana (Glycymeris) Anderson, 1958: 98, pl. 74, figs. 9. 2a; CASG 61874.01. Glycymeris veatchii (Gabb,
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1864), Turonian to late Squires (2010: 902).
pacifica (Opisoma) Anderson, 1958: 122, pl. 26, figs. 5-6; CASG 29118.03. Opis (Hesperopis) triangulata (Cooper, 1894), late Campanian through early late Maastrichtian, CA, BX. Squires and Saul (2009: 1329).
pacificus (Inoceramus ) Anderson and Hanna (1935: 29, pl. 10, fig. 4); CASG 1430.01, early Maastrichtian, BX (Miller and Abbott, 1989: 63, fig. 1). Anderson and Hanna’s name is a junior homonym of Inoceramus pacificus Woods (1917: 28, pl. 14, figs. 1, 2) from Upper Cretaceous (lower Santonian) strata on South Island of New Zealand (Crampton et. al, 2000: 322, fig. 12). Anderson and Hanna’s junior homonym is renamed herein as Inoceramus georgeedavisi new species Squires, in honor of the late George Edward Davis, geologist and molluscan paleontologist.
pac ificus ( (Pectunculus) Anderson, 1902: 74, pl. 7, fig. 159; voucher GASG 70504. Glycymeris pacifica ( Usdin, 1902), late Cenomanian to Turonian, OR, CA, BX. Squires (2010: 898).
peninsularis (Corbis) Anderson and Hanna, 1935: 31, pl. 10, fig. 1; UCMP 36119. Calva (Calva) peninsularis (Anderson and Hanna, 1935), latest Campanian and early Maastrichtian, CA, BX. Saul and Popenoe (1992: 17).
roguensis (Avicula) Anderson, 1958: 106, pl. 38, fig. 7; CASG 445.17. Pteria pellucida (Gabb, 1864), Turonian to Santonian, OR, CA. Squires (2014: Dil),
rosarioensis (Opis) Anderson and Hanna, 1935: 31, pl. 10, figs. 2, 3; UCMP 36120. Opis (Hesperopis) rosar- ioensis (Anderson and Hanna, 1935), early middle to middle late Campanian, CA, BX. Squires and Saul (2009: 1322).
shastaensis (Glycymeris) Anderson, 1958: 97, pl. 19, figs. 8-10; CASG 27830.01. Glycymeris veatchii (Gabb, 1864), Turonian to late Campanian, VI, CA, BX. Squires (2010: 902).
taff (“Trigonocallista”) Anderson, 1958: 139; CASG
Base OL, Calva (Egelicalva) taffi (Anderson, 1958),
Ganiheian to Santonian, CA. Saul and Popenoe (1992: 34).
wheelerensis (Trigonia) Anderson, 1958: 116, (unfig.): CASG 10632. Yaadia leana (Gabb, 1877), Coniacian to early Turonian, OR, CA. Saul (1978: 33).
Campanian; VI, CA, BX.
Gastropods
argonautica (Cypraea) Anderson, 1958:177, pl. 21, figs. 4, 4a; CASG 61856.05. Protocypraea argonautica (Anderson, 1958), Turonian, OR, CA. Lorenz (2017: 205).
bellavistana (Acteonina) Anderson, 1958: 157, pl. 29, fig. 2; CASG 1293.01. Paosia californica (Gabb, 1864). late Cenomanian to Turonian, VI, OR, CA, BX. Squires and Saul (2004a: 492).
berryessensis (Acteonina) Anderson, 1958: 157, pl. 29, fig. 3; CASG 3192.01. Paosia californica (Gabb, 1864), late Cenomanian to Turonian, VI, OR, CA, BX. Squires and Saul (2004a: 492).
eed S ypraea) Anderson, 1958: 176, pl. 63, figs. 2, 2b; CASG 31918.02. Protocypraea ERAT eae 1958), Turonian, CA. Lorenz (2017: 205). biconica (Gosavia) Anderson, 1958: 175, pl. 75, figs. 3, 3a; CASG 61935.01. Konistra biconica (Anderson, 1958), Turonian, CA. Saul and Popenoe (1993: 381). colusaensis (Acteonina) Anderson, 1958: 158, pl. 21, fig. 14: CASG 1291.05. Paosia colusaensis (Anderson, 1958), late Albian to early Cenomanian, CA. Squires and Saul (2004a: 491). condoniana (Anchura) Anderson, 1902: 76, pl. 8, fig. 179: CASG 445.30. Anchura (Helicaulax) condoniana Anderson, 1902, Turonian, OR, CA. Saul and Popenoe (1993: 354). crossi (Volutoderma) Anderson, 1958: 174, pl. 16, figs. 3, 3a; CASG 61934.02. Cydas crossi (Anderson, 1958), Turonian, CA. Saul and Popenoe (1993: 362).
frazierensis Acteonella Anderson, 1958: 161, pl. 29, figs. 7,
7a; CASG 28103. Trochactaeon (Trochactaeon) frazierensis (Anderson, 1958), Turonian, CA. Sohl and Kollmann (1985: 79).
garzana (Phasianella) Anderson, 1958: 162, pl. 73, fig. 5; CASG 28311.02. Tylostoma? garzana (Anderson, 1958), late early to early late Maastrichtian, CA. Squires and Saul (2004b: 27).
gualalaensis (Cypraea) Anderson, 1958: 176, pl. 62, figs. 8, 8a; CASG 61918.01. Protocypraea gualalaensis (Anderson, 1958), early Maastrichtian, CA. Lorenz (2017: 205).
jacksonensis Volutoderma Anderson, 1958: 174, pl. 21, fig.
1; CASG 445.16. Drilluta jacksonensis (Anderson, 1958), Turonian, OR, CA. Saul and Popenoe (1993: 368).
nortonensis Bullina Anderson, 1958: 178, pl. 21, figs. 13, 13a; CASG 61850.03. Ellipsoscapha nortonensis (Anderson, 1958), Campanian, CA. Stecheson (2004: 102).
packardi Acteonella Anderson, 1958: 160, pl. 29, figs. 4, 4a, 4b: not in CASG database. Trochactaeon (Tro- chactaeon) packardi (Anderson, 1958), Turonian. Soh] and Kollmann (1985: 83).
robertiana (Nerinea) Anderson, 1958: 155, pl. 66, fig. 3; UCMP 33954. Turritella chaneyi Merriam, 1941, early Maastrichtian, CA. Saul (1983: 81), Saul and Squires (1998: 465).
roguensis (Acteonina) Anderson, 1958: 158, pl. 30, figs. 5, 5a; CAS 61906.01. Paosia californica (Gabb, 1864), late Cenomanian to Turonian, VI, OR, CA, BX. Squires and Saul (2004a: 492).
rustica (Acteonella) Anderson, 1958: 161, pl. 29, fig 5; CASG_ 33721.01. Trochactaeon (Trochactaeon) frazierensis (Anderson, 1958), Turonian, CA. (Sohl and Kollmann, 1985: 79, 85).
stewarti (Nerinea) Anderson, 1958: 155, pl. 30, figs. 2, 3; CASG 61842.02. A cold-seep epitoniid, ?middle Turonian, CA. Saul and Squires (1998: 465).
ursula (Acteonina) Anderson, 1958: 158, pl. 63, fig. 4 CASG 31210.01. Paosia ursula (Anderson, 1958), Coniacian, CA. Squires and Saul (2004a: 494).
ursulagorda (Acteonina) Anderson, 1958: 159, pl. 63, fig. 5: CASG 31210.02. Paosia ursula (Anderson, 1958). Coniacian, CA. Squires and Saul (2004a: 494).
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siskiyouensis (Gyrodes) Anderson, 1902: 76, pl. 8, figs. 167, 168: CASG 41. Natica conradiana (Gabb, 1864), Turonian, OR, CA. Popenoe et al. (1987: 92).
yrekensis (Acteonina) Anderson, 1958: 157, pl. 29, fig. 2; CASG 61847.01. Paosia californica (Gabb, 1864), late Cenomanian to Turonian, VI, OR, CA, BX. Squires and Saul (2004a: 492).
ARNOLD
Bivalve
stantoni (Mactra) Arnold, 1908: 357, pl. 31, fig. 3; USNM 31001. Cymbophora_ stantoni (Arnold, 1908), Campanian, CA. Saul (1974: 1076).
Gastropod
pescaderoensis (Turritella) Arnold, 1908: 358, pl. 31, fig. 7; CASG 69777. Turritella chicoensis pescaderoensis Arnold, 1908, late Campanian, CA. Saul (1983: 64).
COOPER
Bivalves
triangulata (Corbula) Cooper, 1894: 49, pl. 2 [not 4], fig. 42; lectotype CASG 624. Opis tana! trian- gulata (Cooper, 1894), late Campanian to early Maastrichtian, CA, BX. Squires and Saul (2009: 1329).
bowersiana (Cucullaea) Cooper, 1894:48, pl. 5, figs. 61, 62; CASG 66028.0. Calva (Egelicalva) bowersiana (Cooper, 1894), late early through late Campanian, VI, CA. Saul and Popenoe (1992: 39).
Senta ety
capuloides ( Sieanania) Cooper, 1894: 47, pl. 2, figs. 38 39; CASG 612. Vasculum? capuloides (C Jooper, 1894), late Campanian to possibly early Maas- trichtian, CA. Coan (1981: 160).
cretaceum (Sistrum (Ricinula?)) Cooper, 1896: 330, pl. 47, figs. 1, 2; neotype LACMIP 9997. Perissitys cretacea (Cooper, 1896), Coniacian, CA. Popenoe and Saul (1987: 12).
fairbanksi (Cerithium) Cooper, 1894: 44, pl. 1, fig. 12; not in GASG database. Possibly Anchura, late C: ampanian to possibly early Maastrichtian, CA. Coan (1981: 163).
kempiana ( (Calliostoma) C Jooper, 1894: 46 [pl. 2 [not 3], figs. 33, 34; CASG 610. Calliostoma kempianum Cooper, 1894, late Campanian to possibly early Maastrichtian, CA. Coan (1981: 165).
normalis (Tornatella) Cooper, 1894: 46, pl. 2 [not 3], figs. 36, 37; CASG 625. Eoacteon normalis (Cooper, 1894), late Campanian to possibly early Maas- trichtian, CA. Coan (1981: 168).
DAILEY and POPENOE
Bivalves
apletos (Glycymeris) Daily and Popenoe, 1966: 8, pl. 1 figs. 1, 5, 6; LACMIP 8891. Glycymerita veatchii (Gabb, 1864), middle Turonian to late Campanian, VI, CA. Squires (2010: 902).
pozo (Corbula) Daily and Popenoe, 1966:19, pl. 5, figs. 6-10; LACMIP 8916. Panzacorbula pozo (Daily and Pope- noe, 1966), early Campanian to early late Maastrichtian, CA. Squires and Saul (2004¢: 117).
Gastropod
ainiktos (Pseudoglauconia?) Daily and Popenoe, 1966: 21, pl. 6, figs. 3, 5, 6; LACMIP 8291. Bullamirifica ainiktos (Dailey and Popenoe, 1966), middle through late Campanian, CA, BX. Squires and Saul (2005: 141).
HALL and AMBROSE
Gastropod
branneri (Cerithium) Hall and Ambrose, 1916: 70, unfigured; CASG 69804. Cerithium? teslaensis Hanna, 1924-162, Late Cretaceous, CA. See Wiedey (1929: 25, pl. 1, fig. 6).
McLELLAN
Bivalves
suciensis (Cucullaea) McLellan, 1927: 132, pl. 17, figs. 4-6; UWBM 15010, Glycymerita veatchii (Gabb, 1984), middle Turonian to late Campanian, VI, WA, CA, BX. Squires (2010: 902).
suciensis (Glycymeris ) McLellan, 1927: 131, pl. 17, figs. 7, 8; UWBM_ 15008, Glycymerita veatchii (Gabb, 1864), middle Turonian to late Campanian, VI, WA, CA, BX. Squires (2010: 902).
MERRIAM
Gastropod
tolenasensis (Turritella) Merriam, 1941: 62, pl. 1, figs. 14, 15: UCMP 15328. Turritella hearni Merriam, 1941, Turonian and probably early Coniacian, CA. Squires and Saul (2006a: 54).
MURPHY AND RODDA
Gastropods
allisoni (Gyrodes) Murphy and Rodda, 1960:842, pl. 101, figs. 18-20; LACMIP 9828. Natica? allisoni (Murphy and Rodda, 1960), Cenomanian, OR, CA. Popenoe et al. (1987: 92).
greeni (Gyrodes) Murphy and Rodda, 1960:543, pl. 101, figs. 27-29; LACMIP 9830. Gyrodes (Sohlella?) greeni Murphy and Rodda, 1960, Cenomanian, CA. Popenoe et al. (1987: 79).
stewarti (Sollariella) [sic] Murphy and Rodda, 1960:839, pl. 103, figs. 4, 5; LACMIP 9821. Igonoia stewarti (Murphy and Rodda, 1960), late Cenomanian, CA. Squires (201la: 142).
PACKARD Bivalves
alisoensis (Tellina) Packard (1922: 426, pl. 33, fig. 3); UCMP. 12309. Laternula? alisoensis (Packard,
R.L. Squires, 2018
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1922), middle Campanian, CA. Elder and Saul (1993: pole 2, ress, IL)
angulata (Meretrix) Packard (1922: 425, pl. 33, fig. UCMP 12307. Etea angulata (Packard, 1 Campanian, CA. Saul (1982: 73).
californica (Exogyra) Packard, 1922: 421, pl. 27, fig. 5; UCMP. 12320. Costagyra californica (Packard, 1922), Turonian, CA. Squires (2017: 39).
californiana (Trigonia) Packard, 1921: 17, pl. 2, fig. 2; UO 6 (lost, but CASG 5658 is a plaster cast of it). Popenoella californiana (Packard, 1921), Turonian, CA. Saul (1978: 39), Cooper and Leanza (2017: 329).
chicoensis (Spisula) Packard, 1916: 300, pl. 27, figs. 6, 7; UCMP 12205. Willimactra (Petromactra) truncata (Gabb, 1864), early to late Campanian, VI, CA. Saul (G72 LS).
cordiformis (Cucullaea?) Packard, 1922: 417, pl. 24, fig. 1; UCMP 12311. Cucullaea (Idonearca) Condiformis Packard, 1922, late Campanian, CA. Saul (1982: 68).
coronaensis (Cardium) Packard, 1922: 424, pl. 30, fig. 2: UCMP 12281. Pachycardium coronaense (Packard, 1922), Turonian, CA. Popenoe (1937: 388).
crescentica (Ostrea) Packard, 1922: 420, pl. 26, fig. 3 [not fig. 4]; UCMP 12318. Curvostrea crescentica (Packard, 1922), Turonian, CA. Squires (2017: 29).
demessa haidana var. (Acila (Truncacila)) Packard in Schenck, 1936: 50, pl. 2, figs. 3, 4, 6, 10; CASG 69081. Acila (Truncacila) haidana Packard in Schenck, latest Albian (probably) to early Turonian, QC, CA. (Squires and Saul, 2006b: 92).
deschutesensis (Trigonia) Packard, 1921: 24, pl. 10, fig. 3; UO 9. Notoscabrotrigonia oregana (Packard). middle Albian?, perhaps middle or late Cenomanian, OR. Jones (1960: 437), Cooper (2015: 25).
evansana var. oregona (Trigonia) Packard, 1921: 26, pl. 9, fig. 7; UO 4. Notoscarbrotrigona oregano (Packard), middle Albian to perhaps middle or late Cenomanian, OR. Jones (1960: 437), Cooper (2015: 25).
fitchi (Trigonia) Packard, 1921: 20, pl. 6, fig. 3; pl. 7, fig. 2; UO 26859. Louella fitchi ( (Packard, i991), early ie middle Turonian, OR, CA. Saul (1978: 53), Cooper and Leanza (2017: 330).
hardingensis (Homomya) Packard, 1922: 423, pl. 32, figs. la, Lb: UCMP 12291. Liopistha (Psilomya) hardingensis (Packard, 1922), Turonian, CA. Popenoe (1937: 383).
inezana (Trigonia) Packard, 1921: 27, pl. §, figs. la, b; pl. 9, fig. 1, pl. 10, fig. 1; UCMP 31464. Notoscabrotrigona evansana (Meek, 1858), Coniacian through Campa- nian, VI, OR, CA. Jones (1960: 436), Cooper (2015: 25).
inornata (Exogyra) Packard, 1922: 420, al, Bi, img, lly UCMP 12284. Phygraea inornata (Packard, 1922), Turonian, CA. Squires (2017: 46).
lapidis (Astarte) Packard, 1922: 423, pl. 30, figs. 4a, 4b; UCMP. 12285. Eriphyla lapidis (Packard, 1922), Campanian? CA. Popenoe (1937: 387).
nitida (Meretrix) var. major Packard, 1922: 425, pl. 33, fig. 2; UCMP 12279. Calva (Egelicalva) Doemana (Cooper, 1894), late early through late Campanian, WA, CA. Saul and Popenoe (1992: 39).
5); 922).
ovoides (Astarte) Packard, 1922, 424: pl. 30, fig. 1; UCMP 12280. Eriphyla ovoides (Packard, 1922), late Turonian to Coniacian’, CA. Popenoe (1937: 386), Saul (1982: 72).
striatus (Spondylus) Packard, 1922: 422, pl. 29; UCMP 12276, [a junior homonym renamed as Spondylus fucatus Hanna, 1924: 181], late C Jampanian to early Maastrichtian, CA. Elder (1991: E10).
subnodosa (Lima) Packard, 1922:421, pl. 28; UCMP 12275. Spondylus subnodosus (Packard, 1922), late Campanian to early Maastrichtian, CA, BX? Elder (1991: E10).
sulcata (Astarte?) Packard, 1922: 424, pl. 33, fig. 6; UCMP 12305, [a junior homonym renamed as Astarte? acerba Hanna, 1924: 157 and as Astarte? earllergyi Anderson, 1958: 121: now Alleinacin acerba (Hanna, 1924)]. Turonian, CA. Squires and Ritterbush (1981: 896).
taxidonta (Ostrea) Packard, 1922: 420, pl. 26, fig. 2: UCMP. 12317. Acutostrea taxidonta (Packard, 1922), Turonian, CA. Squires (2017: 32).
Gastropods
californica (Gyrodes) Packard, 1922: 429, pl. 35, figs. 2a, 2b; UCMP 12300. Euspira Shuntardiana (Gabb, 1864), late Campanian, CA. Pope noe (1937: 398).
californiensis (Lysis) Packard, 1922: 431, pl. 37, figs. 2 UCMP 12287. Lysis suciensis (Whiteaves, eieza) late early Campanian to early Maastrichtian, VI, WA, CA, BX. Saul and Squires (2008b: 128).
dubius (Siphonalia) Packard, 1922: 431, pl. 35, fig. 5; UCMP. 12304. Saturnus dubius (Packard, 1922), Turonian, CA. Saul and Popenoe (1993: 367).
nodosa (Alaria) Packard, 1922: 430, pl. 26, figs. 5a, 5b; UCMP. 12297. Latiala nodosa (Packard, 1922), Turonian, CA. Saul (1998: 132).
pseudoalveata (Amauropsis) Packard, 1922: 429, not pl. 35, figs. la, 1b, 3; UCMP 12301. Ampullina packardi Popenoe, 1937, early to middle Campanian, CA. Popenoe (1937: 399), Stecheson (2004).
suciaensis (Cerithium?) Packard, 1922: 430, pl. 35, fig. 4 UCMP. 12303. Zebalia suciaensis (Packard, 1922), middle Campanian, WA, CA. Squires and Saul (2003a: 433).
vetus (Aporrhais) Packard, 1922: 431, pl. 36, fig. 1; UCMP 12298. Alarimella veta (Packard, 1922), Turonian, CA. Saul (1998: 134).
tumida (Bullaria) Packard, 1922: 433, pl. 37, fig. 4, UCMP 12289, [a junior homonym renamed as Bullaria obtenta Hanna, 1924: 159], Turonian, CA. Saul (1982: 72).
WARING
Bivalves chicoensis (Isocardia) Waring, 1917: 62, pl. 8, fig. 3; LSJU 399. pace Clisocolus cordatus Whiteaves, 1879: 157, middle Campanian, CA. Popenoe (1937: 390); Smith, 1978: 332). cordata (Macrocallista) Waring, 1917: 62, pl. 8, fig. 1; not in CASG database. Paraesa? lens (Gabb, 1864), Campanian, VI, WA, CA. Saul (1993: 976).
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cowpert ( (Pecten (Propeamusium)) [sic] Waring, 1917: 63, pl. 7, figs. 1, 2; ;: syntypes CASG 61644. Propeamussium (Parpanrussiiin) cowperi (Waring, 1917), late middle Campanian, CA. Sundberg (1989: 61-62). A “variety” of cowperi occurs in a Paleocene wood-fall (chemosynthetic) deposit in northern Japan (Amano et al., 2018: 636).
triangulatus (Crassatellites) Waring, 1917: 59, pl. 9, fig. 1; syntype CASG 397. Cymbophora _ triangulata (Waring, 1917), late Campanian, CA. Saul (1974: 1083).
Gastropods
compressus (Gyrodes) Waring, 1917: 67, pl. 9, fig. 6; CASG 61958.01. Euspira compressa (Waring, 1917), middle Campanian, CA. Stecheson (2004: 64).
crassa (Cancellaria) Waring, 1917: 66, pl. 9, fig. 5; CASG 61644.06. [a junior orasranaa renamed as Gincellann simiana Hanna, 1924: 160], probably middle Cam- panian, CA. No age update found.
plicata (Turris) Waring, 1917: 68, pl. 9, fig. 3; not in CASG database, [a junior homonym renamed as Surcula binotata Hanna, 1924: 181], Late Cretaceous un- differentiated, CA. No age update found.
rotundus (Pugnellus ) Waring, 1917: 67, pl. 9, fig. 10; LS]U 402. Lispodesthes rotundus (Waring, 1917), middle Campanian to early Maastrichtian, CA. Dailey and Popenoe (1966: 22), Stecheson (2004: 43).
templetoni (Solariaxis) Waring, 1917: 68, pl. 9, fig. 22; not in CASG database. Atira ornatissima (Gabb, 1864), latest Santonian to late Campanian, VI, CA, BX. Squires (2010: 1025).
WHITE
Gastropods
californiensis (Nerita) Orcutt, 1887 (based on White’s, 1885: 12, pl. 5, figs. 7, 8 of Nerita sp.); lectotype USNM 1341 la. Nerita (Bajanerita) californiensis (Orcutt, 1887), late Campanian to early Maastrichtian, BX. Squires (1993: 1085).
condoni (Trophon) White, 1889: 21, pl. 3, figs. 4, 5; syntypes USNM = 20122. Praesargana condoni (White, 1889), Turonian, CA. Saul and Popenoe (1993: 360).
dilleri (Scobinella) White, 1889: 25, pl. 4, figs. 1-3; syn- types USNM 20123. Carota dilleri (White, 1889). Turonian, VI, OR, CA, BX. Saul and Popenoe (1993: 374).
dowelli (Gyrodes) White, 1889: 19, pl. 3, figs. 8, 9; USNM 20126. Gyrodes (Gyrodes) dowelli White, 1889, Turonian, VI, OR, CA. Popenoe et al. (1987: 75).
euryostomous (Trochus (Oxystele)) White, 1885: 12, pl. 5, figs. 9-11; not in USNM online database. Homalo- poma euryostoma (White, 1885), early Campanian to early Maastrichtian, BX. Kiel and Aranda-Manteca (2002: 29).
gabbi (Fulguraria) White, 1889: 23, pl. 3, fig. 1; lec- totype USNM 20112. Volutoderma averillii (Gabb,
1864), early Campanian, VI, CA. Saul and Squires (2008a: 226).
hilgardi (Fulgur) White, 1889: 22, pl. 3, figs. 2, 3; USNM 20117. Pentzia hilgardi (White, 1899). early Cam- panian to early Maastrichtian, WA, CA, BX. Squires and Saul (2003b: 57): Stecheson (2004: 83).
marcidulus (Faunus) White, 1889: 20, pl. 4, figs. 12, 13; USNM 20127. Liocium marcidulum (White, 1889), early Campanian, CA. Squires and Saul (2003c: 149).
nexilia (Ceratia) White, 1889: 21, pl. 3, figs. 13, 14; lec- totype USNM 20119a. Acirsa nexilia (White, 1889), early to late middle Campanian, WA, CA. Squires and Saul (2003a: 39).
obstricta (Stomatia) White, 1889: 18, pl. 4, figs. 10, 11; USNM 20124. Ariadnaria obstricta (White, 1889), late Coniacian? to Santonian, CA. Saul and Squires (2008b: 122).
obtusa (Mesalia) White, 1889: 20, pl. 4, figs. 6, 7]; lec- totype USNM 20116a. Acirsa obtusa (White, 1889). late Santonian to early late Campanian, CA. Squires and Saul (2003a: 36).
oppansus (Lysis) White, 1889: 17, pl. 4, figs. 14, 15; USNM 20115. Lysis duplicosta Gabb, 1864, Cam- panian, WA, CA. Saul and Squires (2008b: 125).
pillingi (Cerithium) White, 1885: 13, pl. 5, figs. 3-6; USNM 13408. Echinoaxis pillingi (White, 1885), early Campanian to early Maastrichtian, BX. Kiel and Aranda-Manteca (2002: 33).
totium-sanctorum (Cerithium) White, 1885: 13, pl. 5, figs. 12, 13; USNM 13409. Tympanotonos (Exechocirsus) totiumsanctorus (White, 1885), early Campanian to early Maastrichtian, BX. Kiel and Aranda-Manteca (2002: 34).
wallaense (Solarium) White, 1885: 14, pl. 5, figs. 1, 2; USNM_ 13412 Trochacanthus wallalense (White, 1885), early Campanian to early Maastrichtian, CA, BX. Squires (201 1b: 3).
WHITEAVES
Bivalves
cretacea (Conchocele) Whiteaves, 1874: 266, plate of fossils (figs. 2, 2a); vouchers CASG 61850.04, 61850.5. Thyasira cretacea (Whiteaves, 1874), exact age unknown, VI. Whiteaves (1903: 383).
cumshewaensis (Arca (Nemodon)), Whiteaves (1900: 294: illustrated by Whiteaves 1884: 235, pl. 31, figs. 8, 8a, 8b): lectotype GSC 4915 (see Smith, 1978: 335). Navonavis cusmshewaensis (Whiteaves, 1900), late Santonian, QC. Haggart and Higgs (1989: 61).
meekana (Te Mina) Whiteaves, 1874: 268, plate of fossils a6 6); GSC 5730. Paraesa? lens (Gabb, 1864),
Campanian, V 7 WA, CA. Saul (1993: 976).
nanaimoensis (Tellina) Whiteaves, 1903: 376, pl. 46, fig. 3; GSC 5729. Willimactra (Petromactra) truncata (Gabb, 1864), early to late Campanian, VI, CA. Saul (1973: 23).
R.L. Squires, 2018
Page 97
suciense (Laevicardium) Whiteaves, 1879: 154, pl. 18, fig. 2; GSC 5713. Cymbophora suciensis (Whiteaves, 1879), Campanian, WA, CA. Saul (1974: 1079) used an unnecessary emendation of Whiteaves’ name "suciense," instead of using the name "suciensis."
suciensis (Linearia) Whiteaves, 1879: 146, pl. 17, fig. 12; GSC 5824 (Bolten, 1992: 235). No new name found, hinge characters are unknown, and the only known ena n has been lost (Squires and Goede srt, 1994: 262); thus, this bivalve is a nomen dubium, early middle Campanian, WA. Squires and Graham (2014: fig. 2).
suciensis (Teredo) Whiteaves, 1879: 135, pl. 17, figs. 1, la; syntypes GSC 5752, 5752 a-d (Bolton, 1992: 239). No new name found, but this fossil cannot be assigned to “Teredo,” whose certain identification requires soft-part morphology, early late Campanian, VI. Squires and Graham (2014: fig. 2).
vancouverensis (Opis) Whiteaves, 1879: 158, pl. 18, figs. 4, 4a; GSC 5691. Opis (Hesperopis) vancouverensis Whiteaves, 1879, middle to late Campanian, VI. Squires and Saul (2009: 1328).
Gastropods
canadensis (Gyrodes) conradiana? var. Whiteaves, 1903: 365, unfigured; GSC 5777. Gyrodes (Sohlella) can- adensis Whiteaves, 1903, Santonian to earliest Campanian, VI. Popenoe et al. (1987: 85).
dakotensis vancouverensis var. (Serrifusus) Whiteaves 1879: 119, pl. 15, fig. 6. GSC 5794 (Bolton: 1965:
74). Serrifusus vancouverensis Whiteaves, 1879.
Boeebly early Maastrichtian, VI. Anderson (1958: 171).
harveyi (Cerithium) Whiteaves, 1903: 362, pl. 43, fig. 7; syntypes 5933, 5933a (Bolton, 1965: 11). Alamirifica? oe (Whiteaves, 1903), Coniacian to early Campanian, VI. Saul and Squires (2003: 448).
intermedium (Mesotoma?) Whiteaves, 1903:360, pl. 43, fig. 4; syntypes 5956, 5956 a-d (Bolton, 1965: 44). Acirsa nexilia (White, 1889), early Campanian to late middle Campanian, WA. Squires and Saul (2003a: 39).
lallierianum suciense var. (Cerithium) Whiteaves, 1879: 122, pl. 15, figs. 10, 10a; lectotype GSC 5764b. Bel- liscala suciense (Whiteaves, 1903), middle to early late Campanian, WA. Squires and Saul (2003a: 33).
newcombii (Mesostoma?) Whiteaves, 1903: 361, pl. 43, fig. 5; GSC 5298 (Bolton, 1965: 45). Confusiscala new- combii (Whiteaves, 1903), latest Santonian to middle Campanian, WA. Squires and Saul (2003a: 40).
nodulosa (Fasciolaria) Whiteaves, 1874: 268, figs. 7, 7a (not 7b); lectotype GSC 5766. Forsia popenoei Saul, 1988. Late early to middle Campanian, VI, CA. Saul (1988: 10), Squires and Graham (2014: fig. 2).
occidentalis radiatula? var. (Solariella) Whiteaves, 1903: 368, pl. 45, figs. 5, 5a; GSC 5918. Igonoia occidentalis (Whiteaves, 1903), Santonian. Squires (201 1a: 144).
suciense (Mesostoma) Whiteaves, 1903: 359, pl. 44, fig. q lectotype GS 5764b. Belliscala suciense (Whiteaves, 1879), middle Campanian to early late Campanian, WA. Squires and Saul (2003a: 33).
suciensis (Cypraea) Whiteaves, 1895: 127, pl. 3, fig. 5; GSC 5937. Palaeocypraea suciensis (Whiteaves, 1895),
early middle Squires and Graham (
Sepa WA. Groves (1990: 275): 2014: fig. 2); Lorenz, 2017: 210).
suciensis (Surcula) Whiteaves, 1879: 115, pl. 15, figs. 1, la
syntypes GSC 5784, 5784 a—b (Bolton, 1965: 79). Amuletum? (Lutema)? suciensis (Whiteaves, 1879), early middle Campanian, WA. Erickson (1974: 223), Squires and Graham (2014: fig. 2).
suciensis carinifera var. (Stomatia) Whiteaves, 1879: 128,
pl. 16, fig. 4; lectotype GSC 5771. Lysis suciensis (Whiteaves, 1879), late early Campanian to early Maastrichtian, VI, WA, CA, BX (including Baja Sur). Saul and Squires (2008b: 128).
suciensis carinifera var. (Stomatia) Whiteaves, 1879: 128,
pl. 16, fig. 5; lectotype UCMP 11975. Lysis duplicosta Gabb, 1864, early to late Campanian, WA, no. CA. Saul and Squires (2008b: 125).
tenuis nanaimoensis var. (Potamides) Whiteaves, 1879: 121, pl. 15, figs. 9, 9a; lectotype GSC 5763a. Anchura nanaimoensis (Whiteaves, 1879), middle to late Campanian, VI. Elder and Saul (1996: 390).
ACKNOWLEDGMENTS
Peter Roopnarine and Christine Garcia provided CASG type-specimen numbers. Lindsey T. Groves (LACM,
Malacology Department) provided detailed information. Lorenz (201 7), provided some type-specimen numbers, critically reviewed the manuscript, and gave valuable comments about it.
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THE NAUTILUS 132(3-4):101-112, 2018
Page 10]
Anatomy of Engoniophos unicinctus from Isla Margarita, Venezuela (Gastropoda: Caenogastropoda: Nassariidae), with a discussion on
the buccinid-nassariid relationship
Daniel Abbate
Luiz Ricardo L. Simone
Museu de Zoologia da Universidade de Sao Paulo Sao Paulo, (4299- 970 BRAZIL danimalacologia@gmail.com
Irsimone@usp.br
Daniel C. Cavallari
Centro para Documentagao da Biodiversidade
Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto Universidade de Sao Paulo
Ribeirado Preto, BRAZIL
decavallari@usp.br
ABSTRACT
The taxonomic position of Engoniophos unicinctus (Say, 1826) in the family Nassariidae is comfirmed based on morphological results. Our detailed anatomical analysis reveals some disparities with typical nassariids, e.g., having a non-serrated operculum,
a simpler and proportionally smaller head, and the absence of
epipodial tentacles. However, the similarities in shell and radula
are present and are easily noticeable, which corroborates the taxonomic data and the positioning of E. unicinctus within the Nassariidae.
Additional Keywords: Anatomy, Buccinidae, Neogastropoda, Taxonomy
INTRODUCTION
The families Nassariidae and Buccinidae are among the most diverse within the Caenogastropoda. They occur from the poles to the equator, inhabiting a wide varie ty of environments, from shallow water to abyssal ocean depths (Harasewych, 1998; the families are small- to medium-sized (20-250 mm), with ovate to fusiform, weakly shouldered shells, an open siphonal canal, columella lacking plicae, and axial sculpture usually consisting of broad aie limited to early whorls. These families ewe a rich fossil record dating from the lower Cretaceous (Taylor and Morris, 1980), together with Fasciolariidae, Melongenidae, and Cancellariidae, while other neogastropod families appeared between the late Cretaceous and early Paleocene (Tracey et al., 1993). The systematics of both families are poorly understood, and there is no consensus as to their limits and their relationships (Simone, 2011). Ponder (1974) considered the Buccinidae closely related to the Nassariidae, Fas-
ciolariidae, and Melongenidae, based on the absence of
accessory salivary lands and an anal gland. This close affinity was emphasized by Kantor (1996). based on
Hayashi, 2004). Members of
shared characteristics such as a long or very long pro- boscis, the loss of glandular dorsal galas. and a te ages ncy toward the reduction of the gland of Leiblein. The mo- lecular systematics work by Galindle et al. (2016) provided a hypothesis for the sy stematic relationships within the Nassariidae.
Buccinids are usually distinguished from the nassariids by having a radula with three or more cusps on the central teeth, hie ral teeth with two large flanking cusps (Golikov, 1980; Kantor, 1990a), and zero to five intermediate cusps (Harasewych, 1998). These radular distinctions, however, have not been observed in Engoniophos unicinctus (Say, 1825), which shows shell and radula greatly similar to those of the Nassariidae (Bandel, 1984). This led some authors to suggest its relocation to Nassariidae, and not in Buccinidae as it is traditionally allocated (Bandel, 1984; Faber, 2004).
The present paper is part of a larger project aiming at an improvement of the taxonomic knowledge on the Bucci- noidea. Focusing in a problematic species with a clear duality of familiar attribution, we intend to discuss its morpho- anatomical attributes which has been published on nas- sariids and buccinids, and provide new data on the species.
MATERIALS AND METHODS
Specimens examined in this study are deposited in the collection of the Museu de Zoologia da Universidade de Sao Paulo (MZSP). A list of examined specimens is given after the species description. Shells were broken prior to soft-tissue extraction. Specimens were immerse din 70% ethanol and dissected by standard techniques under a stereomicroscope. The terminologies of Amold (1965) and Simone (2011) were employed for shell cone TS, and Diver's (1931) whorl-counting method was applied. Details of the radula were examined under the scanning electron microscope (SEM) at the MZSP. The synonymic list presented herein is restricted to taxonomic and oc- currence papers.
Page 102
Abbreviations used in anatomical drawings: aa, anterior
aorta; ab, albumen gland; ae, anterior oesophagus; ag, albumen gland; an, anus; au, auricle; bg, buccal ganglia; br, subradular membrane; ce, cerebral ganglion; eg, cement gland; em, columellar muscle; co, cement gland orifice; ev, ctenidial vein; df, dorsal fold of buccal mass: dg, digestive gland; dl, gland of Leiblein duct; ep, pos- terior oesophagus; es, oesophagus; fp, female pore; fs, stomach folds; ft, foot: ge, suboesophageal ganglion; gi, gill; hg, hypobranchial gland; kd, kidney, ml-ml11. odontophore muscles: il, mantle border; me, medium oesophagus; mj, julgal muscle; mo, mouth; mt, mantle; ne, nephrostome; ng, nephridial gland; nr, nerve ring; oe, odontophoral cartilage; od, odontophore; og, osphradium ganglion; os, osphradium; ot, oral tube; ov, pallial oviduct; oy, Ovary; pa, posterior aorta; pb, proboscis; pd, penial duct; pe, penis; pf, propodial expansion; pg, pedal gland; po, palial oviduct; pp, penial papilla; pr, propodium: pu, pleural ganglia; ra, radula; rm, retractor muscle of pro- boscis; rm, radial ar nucleus; rs, radular sac; rt, rectum; sd, salivary duct; sf, siphon; sg, salivary gland; so, salivary duct orifice; su, suboesophageal ganglia: te, tentacle; va, vaginal atrium; vd, vas de ses -rens: ve, ventricle: vo, visceral oviduct; ye, eye.
RESULTS
Family Nassariidae Iredale, 1916 Subfamily Photinae Gray, 1857
Genus Engoniophos Woodring, 1928 1873 by original
Type Species: Phos erectus Guppy, designation. Miocene, Jamaica.
Engoniophos unicinctus (Say, 1826) (Figures 1—25)
Nassa unicincta Say, 1826: 211-212 (pl. 57, figs 1, la Tryon, 1873: 35 (fig. 55), 1882: 245.
Nassa pallida Powys, 1835: 96.
Nassa guadelupensis Petit de la Saussaye, 1852: 171 (pl. 2 figs & 4).
Strongylocera textilina Morch, 1852: 80 (Lister, pl. 965, fig. 20).
Phos guadalupensis |sic]: Arango, 1880: 201.
Phos guadeloupensis [sic]: Tryon, 1881: 219 (pl. 83, figs 512, 520).
Nassa textilina: Tryon, 1882: 243.
Phos unicinctus: Dall, 1889: 178; Gardner, 1926: 460.
Strongylocera unicincta: Dall and Simpson, 1901: 402.
Engoniophos guadelupensis: Warmke and Abbott, 1962: 115 (pl. 21, fig. F); Humfrey, 1975: 150; Bandel, 1976: 99 (fig. 3a—b).
Engoniophos unicinctus: Rehder, 1962: 131; Warmke and
Abbott, 1962: 116 (pl. 21, fig. J); Work, 1969: 672; Abbott,
1974: 220 (fig. 2428): Wolves andl Volzes, 1983: 26 (pl. 14,
fig. 21); Bande 1, 1984: 142 (fig. 235; pl. 16, fig. 6); Jong and
THE NAUTILUS, Vol. 132, No. 3-4
Coomans, 1988: 83 (pl. 38, fig. 455); Diaz and Puyana, 1994: 187; Ramos and Role. 1994: 102: Faber, 2004: 8: Buitrago et al., 2006: 639; Cruz and Gandara, 2006: 132; Riemivindle z and Jiménez, 2007: 5; Reyes et al., 2007: 384; Rosenberg et al., 2009: 650.
Pallacera unicincta: Macsotay and Campos, 2001: 88. Type Locality: Coast of South Carolina (Say, 1826 in error; see Woodring, 1964: 269).
Distribution: Lower Caribbean to Venezuela (Macsotay and Campos, 2001).
Description: SHELL (Figures 1, 3, 4). Fusiform, twice as long as wide, with 7-8 convex whorls, light gray to cream. Protoconch wide, smooth, dome- shaped, white, with three whorls; transition indistinct. Teleoconch sculpture consisting of 5—S thin spiral cords along entire surface of all whorls (twice as many on body volharal)s space between cords equals three to four times their width, becoming more closely spaced near suture; spiral cords crossed Ib wide axial ribs (width ~1/10 of whorl width) bearing rounded nodules at middle level on earlier whorls, and on upper portion of body whorl; axial ribs becoming less prominent toward suture and basal portion of body whorl; depression between ribs as wide as ribs. Spire angle Oo. Aperture elliptical, twice as long as wide, white, glossy; length ~1/2 of shell length. Siphonal canal short, broad, dorsally recurved. Armall notch present, but not well- marked. Outer lip thick, lirate, with thickened outer edge. Columella straight, without folds.
Heap-roor (Figures 8, 9, 11, 13). Head protruded, ten- tacles elongate d and narrow, twice as long as head; eyes located on “oral protuberances at mid level of tentacle, clearly separating basal broad and distal narrow portions. Rhynchostome as transverse slit, located in middle region of ventral surface of head (Figure 9, 15). Foot large, occupying whole body whorl (retracted), bearing pair Oe smal] propodeal expansions; ene gland located in central region of foot (Figure 11), forming groove, extending from dorsum of foot (propodium) to sole. Cement gland of females rounded, located in anterior region of meso- podium, inner space wide, duct long (Figure 11). Cement gland orifice located on median line of anterior sole region (Figure 13). Opercular pad elliptical, ~80% as wide as dorsal surface of foot; attachment with operculum oc- cupying ~70% of foot area. Penis originated on right lateral region, posterior to cephalic base, at level of mantle edge. Columellar muscle wide and broad, 1% whorl long. Haemocoel long and thin, extending dorsally along center of foot and columellar muscle (Figure Qi),
OpeRcULUM (Figure 2). Small, oval, corneous, pale brown, located close to edge of foot; occupying 1/2 of apertural area. Nucleus terminal, inferior. Outer surface with normal concentric oe lines, forming undulations. Scar oval, occupying ~7/3 of inner aries,
D. Abbate et al., 2018 Page 103
007
Figures 1-7. Engoniophos unicinctus, MZSP 77798 8, shell, operculum and radula. 1. Shell, apertural and dorsal views (L = 18.7 mim). 2. Operculum, inner and outer views. Scale bar = 2 mm. 3-4. Protoconch and first teleoconch whorls. 3. Lateral view. Scale bar = 0.5 mm. 4. Apical view. Scale bar = Imm. 5-7. SEMs of radula. 5. Middle portion of radula. Scale bar = 30 xm. 6. Detail of central teeth. Scale bar = 10 wm. 7. Detail of lateral teeth. Scale bar = 10 pm.
Page 104 THE NAUTILUS, Vol. 132, No. 3-4
po rt
Figures 8-14. Engoniophos unicinctus, detailed anatomy. 8. Complete specimen extracted from shell (operculum removed), an- terior-right view. 9. Detail of head, dorsal view. 10. Reno-pericardial region, ventral view, some adjacent structures also shown. 11. Foot of female, sagital section. 12. Pallial cavity roof, ventral view, and coiled visceral mass. 13. Sole of foot, female, showing propodial
expansions and cement gland opening. 14. Pallial cavity roof, transverse section at middle level of osphradium. Scale bars = 2 mm.
D. Abbate et al., 2018
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MANTLE Cavity Orcans (Figures 10, 12, 14). Mantle cavity covering ~1 whorl. Siphon width ~1/2 of mantle cavity
width, i ngth ~2/3 of mantle cavity length. Right base of
siphon high, width ~twice of mantle e dge Satie left base
low. Osphradium long and narrow, elliptical, length ~1/3 of
pallial cavity length, seagthh ~1/6 of its width. Osphradial filaments short, w aks similar to mantle edge. Ctenidial vein (efferent branchial vessel) with uniformly narrow along its
length. Gill elliptic, occupying ~80% of le ngth and wlVa of
width of pallial cavity. Anterior end of gill aun d, ending gradually, inserted directly into pallial cavity. Gill filaments relatively uniform in size along its entire le ngth. Posterior end of gill rounded, located in posterior region of mantle
cavity Rinse to pericardium. Gill filaments height ~1/3 of
pallial cavity height, apex rounded, at middle portion, tilted to right; right and left ec dges of filament straight; space between gill and right pallial organs about twice of gill
width. Hypobranchial gland inconspicuous. Right side of
pallial cavity almost entirely filled by gonoducts. Rectum long and thin, with ~1/2 of p: allial cavity length. Anus sessile, distanes between anus and mantle border 1/3 of total pallial cavity length. Anal gland absent.
ViscERAL Mass (Figures 8, 10, 12). Extending ~3 whorls posteriorly to pallial cavity. Digestive gland Genk beige, occupying ~S0% of visceral mass Palme! encircling stomach. Gonad orange with small black spots, located on columellar surface, posterior to stomach. Seminal vesicle of males located in anterior portion of gonad, ~% of its size. Kidney occupying ~1/3 of visceral mass volume, located on right side of anterior visceral end. Stomach not seen in ale ail.
CIRCULATORY AND EXCRETORY SysTEMS (Figure 10, 12). Reno-pericardial region occupying 1/3 whorl, oval, on right margin of last soho of visceral mass, adjacent to mantle cavity. Pericardium occupying 1/3 of reno-pericardial region, posterior to gill; situated on left anterior margin of wisearall mass. Aunties reniform, anterior to wreralvatelles with three connections: on upper right side with kidney, on anterior right side with gill, and on posterior left side with ventricle. Vemaaalle four times size of auricle, with common
aorta on left posterior margin. Aortas wide, located along
left-posterior region of ee ele Kidney occupying 1/5 of
pallial cavity volume, renal lobe single, solid, with glandular transverse folds along its ventral Moree etrerent renal vessel located at its right portion. Nephridial gland not seen in detail. Nephrostome a small, transverse slit, located in anterior region of membrane between kidney and pallial cavity.
DicEsTIVE SysTeM (Figures 9, 15-21). Mouth longitudinal, narrow, located on center proboscis tip. Proboscis long, straight and thin, occupying ~80% of hemocoel aeeltinnnes not ‘completely retractable. Rhinchodeal wall thin, in- volving ~1/3 of proboscis. Retractor muscles covering ~ 1/3 of proboscis; several thin retractors muscles originated on
dorsal surface of foot, inserted into posterior end surface of
proboscis. Odontophore and buccal mass muscles: mj, thin pairs of perioral muscles connected on both sides, dorsal and ventral, surrounding odontophore cartilages; m1, jugal muscles, several small muscle fibers connecting buccal mass to adjacent inner surface of proboscis; m2, pair of strong retractor muscles of buccal mass, originating on inner surface of proboscis, running along entire odontophore, inserting into anterior region of odontophore cartilages: m2a, pair of retractor muscles of buccal mass, originating on dorsal surface of haemocoel, inserting at end of posterior margin of odontophore cartilages; m2b, ventral single, thin Paeele auxiliary of m2, originating on ventral medial fibers of m2, detaching from it in region just posterior to m6; m3, long, ~80 % of odontophore le ngth, cylindrical muscle forming outer wall of odontophore, eatlh transverse fibers: m4, pairs of strong radular dorsal tensor muscles cove ring almost entire rane of poste nor por- tion of odontophore cartilages, inserting into subradular membrane; m5, pair of sxovaltierey dorsal tensor muscles of radula, originating inside edges of cartilage, adjacent to rasertion of m4: m6, horizontal muscle, thick, connecting ventral edges of cartilages, running almost along its entire length; m8, pair of small elliptical muscles, length ~1/3 of odontophore cartilage length, originating at anterior end of odontophore cartilages, running along ventral surface of odontophore, inserting on anterior ventral surface of cartilages; m11, pair of ventral tensor muscles of radula, elongated, about 1/2 of total odontophore length, originating at ventral-posterior end of cartilages, crossing ventrally entire odontophore, inserting into ventral posterior surface of radula. Additional odontophore structures: br, subradular membrane, thin, translucent, along entire length of radular ribbon, covering inner surface of odontophore cartilages; oc, odontophore cartilages, about 3 times as long as wae inner ventral surface concave, ~1/4 of anterior end fused with each other, ~1/3 of anterior region, concave, involving radular ribbon; rs, radular sac thin-walled, cylindrical, located at posterior end of radula. Radular teeth (Figures 5, 6, 7): rachidian tooth wide, comb-like, occupying about half of radular width; base curved, width ~3% its length; ~9 triangular, sharp pointed cusps of similar size, except for some lateral reduction; lateral tooth hook- like, bicuspid, base broad (equivalent to rachidian base width), obliquely disposed; external lateral cusp widely curved inwards, about as long as base; inner cusp approximately half size of main lateral cusp. Salivary glands (Figure 15) small, located at anterior portion of aeroeoel occupying ~1/8 of haemocoel volume, entirely involving nerve ring, middle esophagus and anterior portion mat proboscis. Salivary ducts very narrow, except for short proximal region running completely attached to anterior esophagus wal] and, more anteriorly, inside dorsal folds of buccal cavity (Figure 16); opening very small (Figure 16: so), in anterior-middle region ae dorsal folds of buccal cavity. Valve of Leiblein (Figures 15, 18: vl) large, about 1/8 of odontophore volume, located in medium esophagus anterior to nervous ring, antenor region w ith transverse white band bearing long cilia, middle and posterior regions white, corresponding to inner gland
Page 106 THE NAUTILUS, Vol. 132, No. 3-4
Figures 15-19. Engoniophos unicinctus. Detailed anatomy. 15. Head and haemocoel, ventral view, foot and columellar muscle removed. 16. Proboscis and anterior esophagus opened longitudinally, showing salivary ducts and their apertures. 17. Extended proboscis opened longitudinally, ventral view, odontophore as in situ. 18. Mid and anterior region of posterior esophagus and associated structures, showing valve and gland of Leiblein. 19. Odontophore, dorsal view, superficial layer of membrane removed. Scale bars = 2 mm.
D. Abbate et al., 2018 Page 107
Figures 20-25. Engoniophos unicinctus. Detailed of anatomy. 20. Odontophore, dorsal view, superficial muscles dissected. 21. Odontophore cartilages, dorsal view, some adjacent muscles shown. 22. Penis, ventral view, penis duct shown by translucency. 23-24. Nerve ring, ventral and dorsal views. 25. Pallial oviduct, ventral view, transversely sectioned at its middle level, some adjacent structures also shown. Scale bars = 2 mm.
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occupying most of inner surface; bypass absent. Gland of
Leiblein narrow and elongated (Figures 15, 18: gl), ~twice as long as middle esophagus, becoming gradually narrower posteriorly; duct of gland of Leiblein straight, long and very narrow, length ~1/2 of medium esophagus length, width ~1/6 of medium esophagus width. Accessory salivary glands absent. Anterior esophagus broader; walls muscular, with several dorsal internal longitudinal folds, occupying entire length of proboscis. Middle esophagus slender, short,
diameter and length approximately half those of anterior esophagus. Posterior esophagus narrow, straight, about as long as anterior esophagus; anterior region broad, differentiation between middle and posterior esophagus clear with a thin duct. Stomach not seen in detail.
REPRODUCTIVE SysTEM, MALE (Figure 22). Vas deferens narrow, simple, straight, running along ventral surface of kidney up to pallial cavity. Prostate totally closed
Figures 26-28. Hard structures of some nassarids for comparison. 26. Nassarius albus MZSP: 109759 shell, apertural and dorsal views. 27. Operculum of Nassarius arcularia plicatus MZSP: 99863 outer view, scale bar = 2 mm. 28. Radula of Nassarius arcularia plicatus, SEM, MZSP: 99863, detail of central and lateral teeth. Scale bar = 20 pm.
D. Abbate et al., 2018
Page 109
(tubular), running through right mantle edge, at ~2/3 of total pallial cavity length. Vas deferens anterior to
yrostate, straight, running immersed into integume nt of I
dorsum, next to mantle arias sr and penis base. Penis large, length ~1/2 of total head-foot length, dorso-ve ori flattene -d; base curved, apical region pointed, Penial duct
29 cm
et
Figures 29-31.
straight, running through center of penis, closed (tubular). Penial papilla long, slender, subterminal, located at anterior left region of penis. Penial aperture apical, far from papilla. REPRODUCTIVE SYSTEM, FEMALE (Figures 10, 12, 25). Visceral oviduct narrow, straight, running along ventral surface of
30
Op
Anatomy of Nassarius vibex. 29. Complete specimen extracted from shell ( (operculum removed), right-anterior view.
30. Penis, ventral view, transverse section at indicated levels also shown. 31. Head-foot, dorsal view, showing met :podis al tentacles. Scale
bars = 2 mm.
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kidney in first whorl of visceral mass, located in front of
pallial cavity. Posterior region of pallial oviduct protruding into kidney, occupying almost entire length of pallial cavity and 1/3 of its width. Albumen gland posterior, whitish, thick-walled, occupying ~1/5 of pallial oviduct volume. Capsule gland ~4/5 of oviduct volume, elongated, orange, thick- male d. Vaginal atrium anterior to capsule Bland occupying ~1/7 of oviduct volume: sinuous, walls nck. muscular, female genital pore narrow, protruded, papilla- like, with thick edges. Bursa copulatrix absent.
CreNTRAL Nervous SysTEM (Figures 25, 26). Nerve ring lo- cated in ventral basal proboscis region (Figure 15: nr), occupying ~ 1/12 of haemocoel volume, highly concentrated.
Ganglia mostly fused, hard to distinguish from each other, SOE asymmetrical. Pairs of pleural and cerebral ganglia fully fused with each other. Sub- esophageal ganglion about half of cerebro- pleural ganglion size, shortly
andl broadly connected to right cere beal ganglion. Pair of buccal ganglia small, located close to posterior edge of
cerebral ganglia. Esophageal aperture small, occupying about 1/9 of ventral surface of nerve ring. Statocysts not seen.
Shell Measurements (length < width in mm): MZSP THis YS LO. K 1O¢Ze 175 XK VOr GS IBD) X< Bil 18.7 X 9.5.
Habitat: Muddy and sandy bottoms, intertidal.
Material Examined: MZSP 77798: 26, 22, Venezuela: Isla Margarita; Playa Bella Vista, 10°56’ N, 63°50 W, 3-4 m als »pth ( (Simone col., 02/ii/1995).
DISCUSSION
Engoniophos unicinctus has a clear conchological re- semblance to some representatives of Nassariidae, e.g., Tritia alba (Say, 1826) (Figure 26), in having a small shell with a poorly develope d parietal callus, a high spire, short siphonal canal, weak anal notch, and uniform sculpture throughout the- shell surface. Moreover, there is also
a resemblance in radular configuration. The radula of
E. unicinctus shows lateral reas bearing two similarly sized cusps, lacking the smaller alert FEU teeth present in Buccinum undatum Linnaeus, 1758, type species of Buccinidae (Fretter and Graham, 1962: 171, fig. 105E). On the other hand, the species in the nassariid genus Buccinanops also bears intermediate cusps in lat- eral teeth. It also shows rachidian teeth with a larger number of cusps, as in Nassarius arcularia plicatus
(Roding, 1798), a subspecies of the type species of
Nassariidae (Figure 28). Such characteristics led Faber (2004) to argue that E. unicinctus would be better placed in Nassariidae, following Bandel’s (1984) suggestion, in- stead of being allocated in Buccinidae, as proposed by certain authors in more traditional classifications (e.g., Cernohorsky, 1984; Miloslavich, 1999; Reyes, 2007).
Engoniophos unicinctus presents some anatomical differences from those of typical nassariids, for example, the operculum (Figure 2) lacking ae serrations commonly found in nassariines (Figure 2 27), but these are also absent in dorsanine fiiceinanopanes The head of nassariids is generally well developed (Figure 29), while in E. unicinctus it is proportionally much smaller and sim- pler, with the tentacles inserted directly into the dorsum (Figure 8: te). The penis is short and broad, with a long and slender papilla in E. unicinctus (Figure 22), while in other nassariids it is normally long and slender, lacking papillae (Figure 30). The epipodial tentacles (Figure 31), either singly or in pairs, are a traditional character and a demonic anatomical feature of Nassariidae (Simone and Pastorino, 2014). This feature is not found in E. unicinctus, which only bears a pair of propodeal (anterior) expansions (Figure 13), also ee found in other nassariids, but no tentacles. Last but not least, recent molecular analyses, place E. unicinctus within the Nas- sariidae (Galindo et al., 2016).
The morphological similarities above mentioned common with nassariids and the aforementioned recent findings in the literature so far indicate that E. wnicinctus, in fact, belongs to Nassariidae. This conclusion adds an- atomical argumentation to the already known conchological and anole similarities, and further corroborates the fa- miliar placement, and its positioning within Buccinoidea proposed by Galindo et. al (2016).
ACKNOWLEDGMENTS
The authors are grateful to Lara Guimaraes (MZSP) for helping with the SEM examination. This work was par- tially supported by a doctoral grant by the Conselho
Nacional de Desenvolvimento Games e Tecnolégico (CNPq, proc. No 159448/2012-3).
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THE NAUTILUS 132(3—4):113-116, 2018
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Mericella zhangsupingae, a new cancellariid species from the South China Sea (Gastropoda: Cancellariidae)
Shugian Zhang’
Peng Wei
Laboratory of Marine Organism Taxonomy and Phylogeny Institute of Oceanology, C hinese Academy of Sciences Qingdao 266071, CHINA
And
Center for Ocean Mega-Science, Chinese Academy of Sciences Qingdao, 266071, CHINA
ABSTRACT
A new western Pacific species belonging to the family Can- cellariidae is described from off Dongsha Islands, South China Sea, at bathyal depths of 500-600 m. Mericella shangsupingae new species is most similar to Mericella bozzettii Petit and Harasewych, 1993 in shell size and proportion. However, the new species can be separated from that species in having a larger, depressed protoconch, more convex teleoconch whorl, much weaker sculpture, and more sinuous outer lip. The new species represents the first member of genus Mericella discovered from westerm Pacific.
Additional Keywords: Volutoidea, Dongsha Islands, China, protoconch
INTRODUCTION
The genus Mericella was originally established by Thiele (1 929), as a subgenus of Craccionio, to accommodate a single species previously treated as Cancellaria (Merica) jucunda Thiele, 1925. To date, three species have been recognized in the genus, including Mericella jucunda (Thiele, 1925) fost off Tanzania, Mericella paschalis (Thiele, 1925) from off Tanzania and Mozambique, and
Mericella bozzettii Petit and Harasewych, 1993 from off
Somalia. Olsson and Bayer (1972) proposed the generic name Gerdiella to include three cancellariid species from bathyal depths (516-897 m) of the Florida Straits and the Caribbean Sea. In that publication, they recognized that Gerdiella was related to, and possibly congeneric with, Mericella, but would differ by shell size and “geographical distribution. Verhecken and Bozzetti (2006) compared the two genera based on conchological characters, and found no reason for further generic separation, although recognizing some degree of difference (e.g. spire height, suture form, relatively aperture height). However, a
l ; ~ ae Author for correspondence: zsqtaxon@qdio.ac.cn
significnt distinction between the two genera was over- looked by Verhecken and Bozzetti (2006). Mericella species have a smooth protoconch, while Gerdiella species have axial ribs on the protoconch. In addition, some microscopic spiral threads were observed on the proto- conch of Gerdiella alvesi Lima, Barros, and Petit, 2007. Verhecken (2002) stated that protoconch characters are of no diagnostic importance at the generic level. This opinion was not followed by Lima et al. (2007), who regarded Gerdiella as a distinct genus endemic to the western and southern Atlantic Ocean.
In this paper we describe a western Pacific species belonging to the Cancellariidae and assign it to the genus Meric ila based on its smooth protoconch and geographic proximity to other species of Mericella. The finding ex- tends the distribution of Mericella from East Neen waters to the western Pacific.
MATERIALS AND METHODS
Specimens were trawled by fishermen from bathyal depths (500-600 m) off Dongsha Islands, South China Sea (see Figure 1). Shells were observed using a light microscope and protoconch characters by scanning electron microscope (SEM). All specimens have been deposited at the Marine Biology Museum of Chinese Academy of Sciences (MBMCAS). The following ab- breviations are used in the text: MBM, Marine Biological Museum, Qingdao, China; USNM, National Museum of National History, Smithsonian Institution, Washington, DC.
SYSTEMATICS Family Cancellariidae Forbes and Hanley, 1851 Genus Mericella Thiele, 1929
Type Species: Cancellaria jucunda Thiele, 1925 (off Dar es Salaam, Tanzania).
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Indian Ocean
@ Mericella jucunda (Thiele, 1925) © Mericella paschalis (Thiele, 1925) #F Mericella bozzettii Petit & Harasewych, 1993
¥ Mericella zhangsupingae new species
30 40 50 60
Geographic distribution of known Mericella species.
Figure 1.
Mericella zhangsupingae new species (Figures 1—12)
Description: Shell (Figures 2-12) large for genus, up to 31.5 mm, solid, elongated, with a tall spire, conical, protoconch rounded, aperture narrow. Protoconch (Figures 11-12) with two smooth, glassy whorls, large, maximum diameter nearly 2 mm, indicating plankto- trophic larval development. Transition to teleoconch distinct, marked by numerous weak axial incremental processes. Protoconch forms smooth keel prior to tran- sition to teleoconch. Teleoconch with up to five roundly convex whorls. Suture constricted, narrow, deep. First teleoconch whorl with four weak initial spiral cords, be- coming five on second whorl, eight on penultimate whor! and ca. 30 on body whorl, occasionally with intercalated spiral threads. Axial sculpture of regularly spaced, sinuate, opisthocline ribs, 17 on first teleoconch whorl, 22 on second whorl, forming sharp nodules and cancellated appearance at intersections with spiral cords. Axial ribs and nodules becoming weak and smoother toward pen- ultimate and body whorl. Incremental lines distinct, dense, forming conspicuous axial ridges between spiral interspaces. Varices up to seven in number, very weak on spire whorls, distinct on body whorl. Aperture narrowly
70
sO) 10) 00) 1910) 420) WE
elongate, elliptical, deflected from coiling axis by 18° (in holotype). Outer lip thickened, with about 20 obsolete denticles along entire length of the flaring, strongly sin- uate outer lip; columellar lip with two developed folds. Radula and soft parts unknown.
Type Material: Holotype, MBM286508 (height 31.5 mm; width 13.8 mm); Paratypes 1-2, MBM286509, all from the type locality.
Type Locality: Off Dongsha Islands, South China Sea, 500-600 m.
Etymology: This new species is named after Prof. Suping Zhang in recognition of her contribution to gastropod taxonomy in China.
Comparative Remarks: Based on its elongate shell with a smooth protoconch and a flaring, sinuate outer lip, we assign this new cancellariid species to genus Mericella. Mericella zhangsupingae new species is most similar to Mericella bozzettii Petit and Harasewych, 1993 (Figures 13-15) in shell size and proportion, but differs in having a larger, lower, more globose protoconch, more convex teleoconch whorls, much weaker axial sculpture, and
S. Zhang and P. Wei, 2018 Page 115
Figures 2-15. Shells of Mericella species. 2-12. Mericella zhangsupingae new species. 2-4. Holotype, MBM286508, 31.5 mm. 5-7. Paratype 1, MBM286509, 26.6 mm. 8-10. Paratype 2, MBM286509, 23.2 mm. 11-12. Protoconch of paratype 2. 13-15. Holotype Mericella bozzettii Petit and Harasewych, 1993, USNM 860315, 28.7 mm.
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more sinuous outer lip. In addition, spiral sculpture is dominant in Mericella zhangsupingae, whereas the axial sculpture is dominant in Mericella bozzettii. Mericella zhangsupingae new species can be clearly distinguished from the other two congeners, Mericella jucunda (Thiele, 1925) and Mericella “paschalis | (Thiele, 1925), by its larger, thicker shell with roundly convex tele oconch
whorls. ana a white color. Previously, all members of
genus Mericella were only known from off East Africa (see Figure 1). The finding of Mericella zhangsupingae new species extends the dicen bution of the genus to the western Pacific Ocean.
ACKNOWLEDGMENTS
We would like to express our sincere thanks to Drs. M.G. Harasewych and José H. Leal for valuable comments and meticulous editing. This research was financially sup- ported by the N Jational Natural Science Foundation of China (3] 750002, 41376167).
PIMP RARURE CED
Lima, S.F.B., R.E. Petit, and J. C.N. Barros. 2007. A new species of Gerdiella (Gastropoda: Cancellariidae) from the South Atlantic Ocean off Brazil with discussion of an undescribed species. The Nautilus 121: 99-103. Olsson, A.A. and F.M. Bayer. 1972. Gerdiella, a new genus of deep- water cancellariids. Bulletin of Marine Science 22: 875-880. Petit, R.E. and M.G. Harasewych. 1993. A new Mericella (Mollusca: Gastropoda: Cancellariidae) from northeastem Africa. Pro- ceedings of the Biological Society of Washington 106: 221-224.
Thiele, J. 1925. Gastropoda der Deutschen Tiefsee- -Expedition,
II. Deutsche Tiefsee-Expedition 17: 35-382.
Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.
Gustav Fischer, Jenna 1: 7B.
Verhecken, A. 2002. Atlantic bathyal Cancellariidae (Neogastropoda: Cancellarioidea): additional data, and description of a new species. Journal of Conchology 37: 505-514.
Verhecken, A. and L. Bozzetti. 2006. New data on East-African ee species, and description of a new species of Scalptia (Neogastropoda: Cancellarioidea: Cancellariidae). Gloria Maris 45: 14-25.
THE NAUTILUS 132(3-4):117-123, 2018
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A newly discovered Paleocene species of Boreocomitas (Gastropoda: Pseudomelatomidae) from eastern Hokkaido, Japan, with implications for the biogeography of the Paleocene
Bering Strait
Kazutaka Amano Krzysztof Hryniewicz Department of Geoscience Institute of Paleobiology Joetsu University of Education Polish Academy of Sciences 1Yamayashiki ul. Twarda 51/55
Joetsu 943-8512, JAPAN 00-818 Warszawa, POLAND amano@juen.ac.jp krzyszth@twarda.pan.pl
Robert G. Jenkins
School of Natural System,
College of Science and Engineering Kanazawa Unive rsity, Kanazawa City Ishikawa 920-1151, JAPAN robertgj@staff-kanazawa-u.ac.jp
ABSTRACT
Two species of the gastropod family Pseudomelatomidae, genus Boreocomitas, ee the new species B. inouei, are described from the Paleocene (upper Selandian to lowest Thanetian) Katsuhira Formation in eastern Hokkaido, Japan. These species represent the first Paleocene record of Boreocomitas. The paper discusses also the new recognition of species of Boreocomitas in Paleocene deposits of westerm Greenland and Denmark. Occurrences of some taxa, including this genus, are suggestive of faunal exchanges between the northern Pacific and the North Atlantic/Artic regions from the middle Paleocene (Selandian) to late Eocene. These exchanges could have resulted from direct marine connections between both areas via the Bering Strait.
Additional Keywords: Paleocene, gastropod, Boreocomitas, new species, palaeogeography
INTRODUCTION
Two species of a Paleocene bathyal gastropod genus including a new species are reported from eastern Hlclakeetidlo, Japan. These species are here assigned to the pseudomelatomid gastropod genus Boece The deep-sea gastropod genus Boreocomitas was proposed by Hickman (1976) as an extinct subgenus of Comitas Finlay, 1926, based on specimens from the Cowlitz and Keasey formations in Oregon, western USA. Recent data show that the age of ane. Cowlitz Formation is middle to late Eocene Sand that of the Keasey Formation is late Eocene to earliest Oligocene (Prothero, 2001). Boreocomitas has also been recorded from the upper Eocene Kovachinskaya Formation in western Kamchatka (Gladenkov et al., 1991).
Many well-preserved molluscan fossils have been de- scribed from the upper Selandian to the lowest Thanetian Katsuhira Formation in eastern Hokkaido, northern Japan
(Figure 1; Amano and Jenkins, 2014, 2017; Amano and Oleinik, 2014: Amano et al., 2015a, b, 2016, 2018). These fossils occur mainly in small calcareous concretions (about 30 cm in diameter) with plant debris, considered to be wood-fall communities (Amano et al. ee 2018). From the locality at Katsuhira along the Urahoro River, two species of Boreocomitas including a new one have been recovered. In this paper, we describe these species and discuss their paleobiogeographical significance.
MATERIALS AND METHODS
Three specimens of Boreocomitas inouei new species and one specimen of Boreocomitas species were obtained from carbonate concretions (20 em in diameter) with many bored wood fragments included in dark gray mud- stones of the ravigulbitie Formation. These concretions are exposed along the Urahoro River, 44 m south from the mouth of the Kokatsuhirazawa River, Urahoro Town, eastern Hokkaido (Figure 1; 42°59'10" N, 143°37'38" E). Rocks in the direct proximity to the locality contain dinoflagellate fossils indicating a late Selandian to ear- liest mene ‘tian age (Amano et al., 2018). These species were associated with 22 species of mollusks (Table 1) and one species of echinoid. Among them, Myrtea ezoensis (Nagao, 1938), and Thyasira oliveri Amano and Jenkins, 2018, were proposed as chemosymbiotic bivalves. As mentioned by Amano et al. (2018), the paleobathymetry of the Katsuhira Formation can be estimated as 200 to 500 m in depth. From these occurrences of mollusks, it is possible to consider the two species of Boreocomitas as members of the deep-sea wood-fall community.
All specimens of Boreocomitas are catalogue d in the University Museum of the University of Tokyo ( UMUT). The associated fauna is stored at Joetsu University of Education (JUE). Classification at family level follows
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Okhotsk Sea
\ Hokkaido Island _-
je - Study area —42N
Pacific Ocean
Figure |
the current taxonomy of Conoidea by Bouchet et al. (2011, 2017).
SYSTEMATIC PALEONTOLOGY
Class Gastropoda Cuvier, 1797
Order Neogastropoda Wenz, 1938
Superfamily Conoidea Fleming, 1822 Family Pseudomelatomidae Morrison, 1966
Remarks: Boreocomitas was originally proposed as a subgenus of the genus Comitas Finlay, 1926, and placed in Turriculinae Boxee 1 1942, of Turridae H. Adams and A. Adams, 1853 by Hickman (1976). Family Pseudome- latomidae including the genus Comitas is characterized by a smooth paucispiral protoconch.
Katsuhira
A SS .
ad so°
Sy A / M
Map showing the locality yielding the two species of Boreocomitas discussed.
Genus Boreocomitas Hickman, 1976
Type Species: Comitas (Boreocomitas) oregonensis Hickman, 1976
Remarks: Boreocomitas is characterized by having a medium to moderately large-sized fusiform shell srati axial nodes on the keel and fine spiral threads, which are less prominent on its wide shoulder than anterior ward. Its anal sinus is broad and moderately deep on the shoulder.
According to Hickman (1976), “the protoconch is missing or worn on all specimens examined, but it is appare sntly paucispiral”, The genus Comitas Finlay, 1926, has a more slender shell, a narrower shoulder, a longer anterior canal and weaker nodes on the periphery than in Boreocomitas. It can be judged that these differences are enough to
separate the two taxa as distinct genera. Although the
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Table 1. Mollusks associated species with Boreocomitas inouei new species.
Species
Leionucula yotsukurensis (Hirayama)
Acila (Truncacila) hokkaidoensis (Nagao) Ezonuculana aff. obsoleta Tashiro
Malletia poronaica (Yokoyama) Menneroctenia plena Kalishevich Pseudoneilonella? sp.
Tindaria paleocenica Amano and Jenkins Propeamussium yubarense (Yabe and Nagao) Limaria sp.
Myrtea ezoensis (Nagao)*
Thyasira oliveri Amano and Jenkins*
Astarte sp.
Cidarina? sp.
Naticidae gen. et sp. indet.
Kangilioptera sp.
Urahorosphaera kanekoi Amano and Oleinik Admete katsauhiraensis Amano, Oleonik and Jenkins Acteocina sp.
Retusa sp.
Biplica paleocenica Amano and Jenkins Striodentalium sp.
Laevidentalium sp.
* Chemosymbiotic species.
pseudomelatomid genus Nekewis Stewart, 1927, re- sembles Boreocomitas, Nekewis can be distinguished from Boreocomitas by having a long and straight anterior canal with a fasciole and ahelltoree anal sinus. Wenz (1938) mistakenly considered Nekewis to be a synonym of the raphitomid genus Clinura Bellardi, 1875. Clinura is very similar to Boreocomitas in having biconic general outline with distinct axial nodes on its periphery. However, Cli- nura has a protoconch that is narrowly conical and di- agonally cancellate. Its diagonally cancellate protoconch demonstfate »s that C Pease belongs in Raphitomidae Bellardi, 1875. Clinura also has an anal sinus whose apex is located near the suture on the shoulder slope.
Boreocomitas inouei new species (Figures 2, 3, 4)
Diagnosis: Moderate-sized, short, biconic Boreocomitas species consisting of five and half whorls and at least one smooth paucispiral protoconch. Surface sculptured with 23 spiral cords on base, nine on keel and six above wide shoulder, and fine sinuous growth lines on last whorl. Apex of anal sinus located at midpoint of shoulder.
Description: Shell moderate-sized, attaining 20.4 mm in height, fusiform, consisting of 5.5 teleoconch whorls and one protoconch whorl. Apical angle ranging from 59° to 73°. Protoconch smooth, ee and with large di- ameter (d = 1.6 mm in paratype UMUT CM 32942). Last whorl very large, occupying about 74% of shell height in holotype; spire low; subsutural band very weak; shoulder slope broad and gently concave. Surface of penultimate
whorl sculptured with one fine spiral cord below keel, six
on keel, shoulder slope smooth, without growth ae sand 21 axial nodes on keel: keel of last einan with 22 axial nodes, 23 spiral cords on base, nine on keel, and six very weak fine cords just above keel and five very faint on shoulder below subsutural band: among spiral cords on last whorl, two cords below keel stronger ae other cords. Anal sinus moderately deep, its apex ‘erat d at midpoint of shoulder slope on last whorl and just below subsutural band on penultimate whorl. Aperture pyriform; outer lip very thin; inner lip covered by thin, narrow callus. Anterior
canal short, slightly broken on anteriormost part, but having slight siphonal fasciole.
Type Material: Holotype, UMUT CM 32793 (shell height, 20.4 mm-+: diameter, 13.5 mm): Paratype, UMUT CM 32794 (shell height, 9.0 mm; diameter, 5.9 mm); Paratype, UMUT CM 32942 (diameter, 7.5 mm-+).
Type Locality: Cliff along Urahoro River, 44 m south from mouth of Kokatsuhirazawa River, Urahoro Town, eastern Hokkaido.
Remarks: This is the first record of Boreocomitas in the Paleocene and from the northwestern Pacific region. Boreocomitas inouei new species is similar to Bor- eocomitas biconica (Hickman, 1976) from the middle to upper Eocene Cowlitz Formation in northwestern Ore- gon in having a similar size (shell height of B. biconica, 20.0 mm), a malcvely low spire and a Sane number of nodes on the last whorl (20 in B. biconica). However, the present new species has higher ratio of diameter (D)/shell height (H) than the B. biconica species (D/H = 0.66 for B. inouei: 0.53 in B. biconica). Also, unlike B. biconica, the present species has a spiral cord below the periphery and no beaded subsutural cords.
Some previously identified species belonging to the now invalid genus Pleurotoma Lamarck, 1799. are now allocated to other genera. Among them, Pleurotoma (Pseudotoma) brevior von Koenen, 1885 (p. 35-36, pl. 2, figs. 5a—c), was described from the Selandian of Copen- hagen, Denmark. Later, this species was re-described as Genotia brevior by Ravn (1939, p. 93-94, pl. 4, fig. Lla—b). Judging from the size, outline, sculpture antl smooth protoconch, P. brevior can be confidently allocated to Boreocomitas. Boreocomitas brevior new combination differs from B. inouei new species in having a more slender, larger shell (ca. 30 mm in height), with fewer nodes on the axial keel (18 to 19 in B. brevior) and distinct spiral cords on the shoulder slope, and with more nu- merous protoconch whorls (3.5 in B. brevior).
The genus Clinura Bellardi, 1875, has a similar shell outline to Boreocomitas, with similar, moderately deep anal sinus and many nodes on its keel. Among the species of Clinura, Clinura sp. 1 from the Sonja Lens of the Selandian Agatdal Formation in the western part of Greenland (Nagssuaq) was illustrated by Kollmann and Peel (1983, p. 97-98, fig. 220). This species is very similar to the present new species in having a short biconic outline, apex of the anal sinus located at the midpoint of the broad shoulder slope, and very fine spiral cords above
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Figures 2-5. 32794: Qa.
3a. Aol view; 3b. Abapertural view; :
the keel. This species also has a smooth paucispiral protoconch, which warrants its allocation to Boreocomitas, not in Clinura. This species can be distinguished from B. inouei new species by its larger shell (ca. 40 mm in he ight), less numerous axial nodes, and by h: wing only one spiral cord below the keel on the last whorl.
Etymology: The new species is named for Kiyokazu Inoue (Obihiro), who collected the holotype.
Distribution: Known only from the type locality, in the upper Selandian to lowe st Thanetian Katsuhira Forma- tion, Urahoro Town, eastern Hokkaido.
Boreocomitas species (Figure 5)
Description: Shell small, 13.1 mm in height, fusiform, consisting of 5 teleoconch whorls; protoconch missing. Apical angle 47°. Last whorl large, occupying about 69% of shell he ight; spire high; subsutural band very weak;
shoulder slope broad and gently concave. Surface of
penultimate whorl sculptured with four fine spiral cords below keel, three on keel, shoulder slope smooth, and 21 axial nodes on keel; keel of last whorl with 20 axial nodes, 12 spiral cords on base, three on keel, and no cord on shoulder below subsutural band. Anal sinus moderately deep, its apex located at midpoint of shoulder slope on last whorl and just below subsutural band on penultimate
Boreocomitas species from the Katsuhira Formation. 2-4. Boreocomitas inouei new species. 2, Paratype, UMUT CM Apical view; 2b. Lateral view showing deep anal sinus of growth lines above shoulder slope. 3. Holotype, UMUT CM 32793; 3c. Apical view. 4. Paratype, UMUT CM 32942, Apicz al view. 5. Boreocomitas sp., UMUT CM 32943; 5a. Apertural view; 5b. Side ae Be Apical view.
whorl. Aperture pyriform; outer lip very thin; inner lip covered by thin, narrow callus. Anterior canal short.
Remarks: This species is similar to Boreocomitas inouei new species as above described. However, Boreocomitas species differs from B. inouei by having a slender form, a higher spire, no spiral cord on the shoulder. Bor- eocomitas oregonensis (Hickman, 1976), the type species of the genus, is similar to Boreocomitas species in having a rather higher spire. However, the type species has some distinct spiral threads on its shoulder.
Distribution: Known only from the type locality of Boreocomitas inouei new speices, in the upper Se landian to lowest Thanetian Katsuhira Formation, Urahoro Town, eastern Hokkaido.
DISCUSSION
The described species possibly lived in a wood-fall community of upper bathyal depths. When Hickman (1976) proposed Boreocomitas, this genus appeared to be confined mainly to the middle Eocene to lower Oli- gocene deposits in Oregon. As noted above, Gladenkov et al. (1991) described and illustrated Comitas (Bor- eocomitas) sp. from the upper Eocene Kovachinskaya Formation in western Kamchatka. However, that shell is poorly preserved, with weak nodes on the keel and
kK. Amano et al., 2018
ave 119) Page 12]
a narrow shoulder. It is uncertain whether this species can be classified in Boreocomitas. Our study shows that Boreocomitas dates back to the Selandian and it had a broad distribution including Denmark, westem Greenland and eastern Hokkaido.
The aporrhaid gastropod Kangilioptera Rosenkranz, 1970 shows the same geographical pattern to Bor- eocomitas. Kangilioptera has been recorded from the upper Danian Kangilia Formation in western Greenland, the Selandian Kerteminde Marl in Denmark and the upper Selandian to lowest Thanetian Katsuhira Formation in eastern Hokkaido (Rosenkranz, 1970; Kollmann and Peel, 1983; Amano and Jenkins, 2014; Schnetler and Nielsen, 2018) (Table 2).
The astartid bivalve Astarte paleocenica Amano and Jenkins in Amano et al., 2018 from the Katsuhira For- mation is similar to A. parvula Kalishevich in Kalishevich et al., 1981, from the Danian Krasnoy arskaya Formation in southeastern Sakhalin and A. trigonula Koenen, 1885, from the Selandian Lellinge Gre end (see also Amano et al. 2018: Schnetler, 2001).
The bivalve genus Conchocele Gabb, 1866, has its earliest confirmed occurrences in the latest Cretaceous (Maastrichtian) of Antarctica, the Danian of western Greenland, and the Thanetian of Spitsbergen (Rosenkranz, 1970; Little et al. 2015; Hryniewicz et al., 2016, 2017). In Spitsbergen, the genus survived to the late Eocene-early Oligocene (Thiedig et al., 1980). There are so far no woneiainel records of @onchucele from the Cretaceous and Paleocene of the northern Pacific area. However, some publications suggest it could have been present in the Pacific during the Late Cretaceous. Con- chocele cretacea has been recorded from the Upper Cretaceous deposits from Vancouver Island area (Whiteaves 1874, jo, Wi, figs. 2. Der). Thyasira cretacea (= Conchocele cretacea) has also been recorded from the Coniacian(?) Funks Formation of Sacramento Valley area
in California (Anderson, 1958). Therefore, the absence of Conchocele in the Upper Cretaceous and Paleocene of
northern Pacific area might be just an artifact of the fossil
record. The genus became very common in the northern Pacific region after the Paleocene (Hickman, 2015: Hryniewicz et al., 2017). This distribution pattern shows that Conchocele populations could have interchanged between the northern Pacific and the North Atlantic/ Arctic regions at least by the late Eocene, or probably earlier.
Moreover, the oldest record of the deep-sea arcid bi- valve genus Bentharca Verrill and Bush, 1898, has been found in the upper Selandian to lowest Thanetian Kat- suhira Formation in eastern Hokkaido (Amano et al., 2015). As pointed by Amano et al. (2015), Barbatia (Acar) hennigi Heinberg, 1978, from the Maastrichtian to Danian deposits in Denmark is possibly an ancestor of this genus.
Adding to Boreocomitas, the four deep-sea mollusks above mentioned occur in both northern Pacific and North Atlantic/Arctic, indicating the sea connection be- tween both areas during the Paleocene, although most paleontologists believe that the Arctic was isolated from the Pacific from the Albian to the latest Miocene (Marincovich et al., 1990, 2002: Thiede et al., 1990: Marincovich, 1993: Marincovich and Gladenkov, 1999: Beard and Dawson, 1999). Deep-sea genera commonly have broad geographic distribution (e.g. Amano et al. 2015a, b; Amano and Jenkins 2017), and similarities be- tween the fauna of the Paleocene Katsuhira Formation from Hokkaido and that of the North Atlantic/Arctic
Paleocene mollusk faunas could be partially caused by that. However, an alternative explanation is that the Bering Strait was to some extent open and connected northern Pacific and Arctic oceans, allowing for faunal interchange between both areas during the Selandian to late Eocene time. When Brikiatis (2014) reviewed the early Cenozoic paleogeography around the Arctic region based on the land mammals and flora, he showed no land bridge around the Arctic Ocean during the Selandian in ie Figure 8. Gleason et al. (2009) also reconstructed the early Eocene paleogeographic map without any land bridge surrounding the Arctic Ocean, based on the
Table 2. Taxa of Paleocene faunas shared between the Northwestern Pacific, Arctic and Northem Atlantic regions.
Region NW. Pacific
E. Hokkaido — SE. Sakhalin Ellesmere Is.
Taxa Age L. Selandian— Danian Paleocene E. Thanetian
Bentharca +
Small Astarte + +
Thyasira oF =P
Conchocele
Kangilioptera tr
Drepanocheilus + +
Aporrhaidae
Boreocomitas +
Reference This study Kalishevich Marincovich and
(1981) Zinsmeister
(1991)
Arctic/ N.Atlantic
Spitsbergen Is. W. Greenland
Denmark
Selandian Danian— Danian— Selandian Selandian ata + + + + + + + + tL > +
Rosenkranz (1970), Ravn (1939), Heinberg Kollmann and (1999), Schnetler Peel (1983) vat Nielsen (2018)
Hryniewicz et al. (2016)
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examination of Nd-Sr isotopes in fossil fish debris from Lomonosov Ridge. Although these data support our hy- pothesis on the Paleogene temporary opening of the Bering Strait, more data are needed to eaten it.
ACKNOWLEDGMENTS
We acknowledge Alan Beu (GNS Science) and Geerat I[- Vermeij (UC Davis) for providing comments to an earlier version of the manuscript. We thank Carole S Hickman (UC Berkeley) for her information on the type specimens of Boreocomitas species. We thank Anton E. Oleinik (Florida Atlantic University) and Kai I. Schnetler (Denmark) for their information on fossil species from the Paleogene in Kamchatka and Denmark. We also express many thanks to Kiyokazu Inoue (Obihoro in eastern Hokkaido) who found the holotype and kindly donated Boreocomitas specimen. This study was supporte d_ by a Grant-in-aid for Scientific Research from the Japan Society for Promotion of Science (C, 17K05691, 2017— 2019) to KA and RG]. Financial support to KH was provided by the Polish National Science Centre (NCN) research grant 2014/15/B/ST10/04886 “The influence of Paleocene/Eocene Thermal Maximum on oceanic chemosynthesis-based ecosystems”.
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Three new species of the genus Humboldtiana (Gastropoda: Pulmonata: Humboldtianidae) from Mexico
Omar Mejia’
Benjamin Lopez
Departamento de Zoologia, ENCB-Instituto Politécnico Nacional
Prolongacion de Carpio y Plan de Ayala s/n C.P. 11340 Ciudad de México, MEXICO
José Ma. Reyes-Gomez
Centro de Cultura Ambiental y Biodiversidad, Bolanos, Jalisco, MEXICO
ABSTRACT
Three new species of the genus Humboldtiana are described. The presence of a mantle mottled with dark spots allocates H paquimei from Chihuahua to the subgenus Gymnopallax. On the other hand, the embryonic whorls, a smooth and granular sculpture distributed over the shell surface, allocate H. wixarika from Jalisco and H. aurea from Chihuahua to the subgenus Humboldtiana s.s. The newly described species are distinguishe d from other species of the genus by the combination of die ll and anatomical characters and by their geographic distribution.
Additional Keywords: Taxonomy, land snail, neotropical region
INTRODUCTION
The land snail genus Humboldtiana von thering 1892 (see Pilsbry, 1927) is comprised by more than 50 species patchily distributed from the southwestern United States to central Mexico (Thompson, 2011). With the exception of three “widely distributed species”, H. buffoniana, H. nuevoleonis, and H. durangoensis, most of the species are microendemic and known only from the type locality. The classification followed herein was proposed by Thompson (2011). It recognizes six subgenera based on shell and reproductive anatomy: Polyomphala Thompson and Brewer, 2000; Gymnopallax Thompson, 2006; Clydo- nacme Thompson, 2006; Aglotrochus Thompson, 2006; Oreades Thompson and Brewer, 2000; and Hum- boldtiana Thompson and Brewer, 2000. The subgenus Polyomphala includes species with depressed shells with nodular tubercles and dart bulbs exposed; the subgenus Gymnopallax is composed of species with mottled mantle with dark gray spot, and shell apex usually de- collated; the subge nus Clydonacme includes species with an embryonic shell sculpture consisting of wavy transverse striations and granular sculpture re stricted to the first postembryonic whorl; the subgenus Aglotrochus
ee Author for correspondence: hmejiag@ipn.mx
comprises species that completely lack the granular sculpture; species in the subgenus Oreades do not have the dart apparatus; and the subge snus Humboldtiana is composed of species with e »mbryonic whorls present (not decollated) and a smooth and granular sculpture dis- tributed over the entire shell surface. Despite the presence of a more or less conserved pattern that sug- gests morphostatic radiation (Gittenberger, 1991), it is difficult to find morphological synapomorphies for the entire genus owing to the fact that the granular sculpture is Angst nt in certain species (Thompson and Brewer, 2000; Thompson, 2006). Nevertheless, the molecular phylogeny clearly supports the monophyly of the genera, although not the monophyly of the subgenera (Mejfa and Zuniga, 2007). Despite of this, we manere to the current classification until a new review of the genus is available. Authorities for the new species are as follows: Hum- boldtiana wixarika new species Mejia, Lopez, and Reyes-Gomez; Humboldtiana aurea new species Mejia and Lé6pez; and Humboldtiana paquimei new species Mejia and Lopez.
MATERIALS AND METHODS
De spite being medium- to large-sized land snails, the species of ae genus Humboldtiana are ve ry elusive and hard to collect. In fact, with selected exceptions noted, most of the species previously named have been described based on very few individuals. For this reason, the de- scriptions of new species herein are based on the holotype and one or two paratypes only. Shell description and morphological measurements were performed according to Thompson and Brewer (2000). In all cases, the first measurement (or counts, for the number of whorls) is from the holotype and the measurements in parentheses are from paratypes | and 2, respectively. The types of the newly described species are deposited at Coleccién Nacional de Moluscos (CNMO), Instituto de Biologia, UNAM, Ciudad de México. Color photographs of the currently described species shells are available at https:// osf.io/3cswm.
O. Mejia et al., 2018
SYSTEMATICS Family Humboldtianidae Pilsbry, 1939 Genus Humboldtiana von Ihering, 1892
Humboldtiana wixarika new species Mejia, Lopez, and Reyes-Gomez (Figures 1-4, 13)
Diagnosis: Medium-sized Humboldtiana with pale-brown shell bearing three equal-sized dark brown bands clearly visible on internal shell surface. Presence of short verge composed by four lobes and penis with two thick longi- tudinal fleshy columns distinguish this species from mihee
species in subgenus Humboldtiana, where dart glands are separated from dart sacs by distance equal to or longer than glands length.
Description: SHELL ( Figures 1—4) globose, outer lip not reflected, pale brown, with three chestnut to dark-brown bands of same size, first band often lighter in color toward end of embryonic whorl, 4 whorls (4.1, 4.0). Embryonic shell caramel in color, with 1.7 whorls (1.8, 1.7), first whorl without sculpture, then with almost imperceptible growth lines. Sculpture of rest of shell consisting of white growth lines more evident toward shell aperture and randomly distributed small oval granules absent toward aperture. Umbilicus completely covered by basal portion of
10 mm
4
Figures 1-4. Humboldtiana wixarika new species. Holotype, CNMO 756. 1) Apertural view 2) Apical view 3) Oblique basal view 4)
Detail of embryonic whorls.
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THE NAUTILUS, Vol. 132, No. 3-4
peristome. Parietal callus thin, transparent to white. Shell height 37 mm (36, 38); shell diameter 39 mm (43, 45); aperture height 28 mm (33, 29); aperture diameter 26 mm (25, 29).
REPRODUCTIVE ANATOMY (Figure 13): Penis short 8 mm (6.5, 7.2), bulbous to globose, tapering at distal end, penis lumen with two thick columns, short verge extending to first third of penis and composed of four lobes. Penis retractor muscle 14 mm long (14, 15). Epiphallus long, cylindrical, measuring 38 mm (46, 34). Flagellum short, 51 mm inle ngth (44, 59), near 0.85 times (1.18, 0.67) the combined length of penis plus epiphallus. Atrium short, 1.28 mm (2.27, 1.65). Lower vagina short and slightly shorter than half penis size, 3.76 mm _ (3.76, 3.40), extending to region of dart sacs; four dart sacs approxi- mately of same size: ds] (2.41 mm), ds2 (2.33 mm), ds3 (2.70 mm), and ds4 (2.07 mm) (3.14, 3.02, 4.00, 3.38) (2.53, 2.17, 3.40, 2.56). Median vagina short, bearing four dart glands forming a ring, dart glands reaching a maxi- mum height of 2.91 mm (3.24, 3.31) and separated from dart sacs by distance of 2.55 mm (2.8, 2.11). Spermathecal duct §1 mm length (128, 114), spermatheca with caecum of 11 mm le meth ( (8, 8) adhering to the albumen gland,
elongated and sac-shaped, witty a length of 11 mm (10, 7).
Type Material: Holotype: CNMO 7561; collected 29 August 2017 by José Maria Reyes Gomez; Paratypes: CNMO 7562 (2): same data as the holotype. All from type locality.
Type Locality: Jalisco, km 29 Carretera Bolanos, Tux- pan, 11.1 km northwest of Bolaios, Jalisco, 2454 m al- titude (21°54'04” N, 103°51/33” W). The individuals were collected on the ground in a pine-oak forest.
Distribution: Known only from type locality.
Remarks: The subgenus Humboldtiana comprises a very disparate group of species where the embryonic whorls are smooth (Thompson, 2006). In this subge snus, three groups of species have been recognized, the Hum- boldtiana buffoniana species group in which the dart glands are located just above the dart sacs, the Hum- boldtiana bicincta species group where the dart sac ap- paratus is reduced, and the Humboldtiana texana species group where the dart glands are separated from the dart sacs by a distance equal or higher than the length of the dart sacs and the epiphallic chamber is absent (Thompson and Brewer, 2000). Humboldtiana wixarika new species belongs to the Humboldtiana texana species group. By having a short verge with four digitiform lobes, this species is similar to H. fasciata (Burch and Thompson, 1957) from the state of Hidalgo, Mexico, but in contrast to H. fasciata,
H. wixarika prese Ss two thick columns in the interior of
the penis instead seven longitudinal folds. Besides, this latter feature, the verge, and the median vagina in H. wixarika are smaller than in H. fasciata. On “ihe ether hand, the shape of the penis of H. wixarika is similar to that of H. balanites from the state of Chihuahua, Mexico,
but in this last species, the verge is multilobed and its size is almost the size of the penis, in opposition to H. wixarika, where the verge is short. Additionally, although in both species the atrium is short, it is quite evident in H. wix- arika, while in H. balanites it is barely distinguishable.
Etymology: The specific epithe t, anoun in apposition, is dedicated to the Wixarika people, better known by their Spanish name, Huicholes, a brave people that reject succumbing to the pressures of the modern world.
Humboldtiana aurea new species Mejia and Lopez (Figures 5-8, 14)
Diagnosis: Medium-sized Humboldtiana with thin and fragile shell. The new species is similar to other species in tne. Humboldtiana texana species complex by having protruding dart bulbs at the base of the dart sacs chile differing from the rest of the species in the group by possessing four sub-equal dart sacs and lacking a caecum on the spermathecal duct. The description is based on the holotype and two paratypes.
Description: SHELL (Figures 5-8) small, subglobose and thin, outer lip slightly thickened, pale-brown to golden with three clearly defined dark-brown bands, in certain specimens bands fade toward the aperture, with first and second band of same width, third band narrower, 4.1 whorls (3.2, 3.2). Embryonic shell with 1.2 whorls (1.1, 1.3), paler than rest of shell. First whorl without sculpture, remainder of whorls with few, weak growth lines. Re- mainder of shell sculptured with growth lines and small ovate granules absent on umbilicus. Umbilicus partially cove ted by basal portion of peristome. Callus thin, transpare nt. Shell height 25 mm (25, 23); shell diameter 29 mm (31, 28); aperture height 19 mm (17, 16), aperture diameter 19 mm (18, 18).
RepropuctivE ANaTomy (Figure 14): Penis short and globose 7.61 mm (6.94, 7.98), penis with verge extending els penis length, composed of three skirts of tissue, one of them with three digitiform processes. Penis retractor muscle 5 mm (5.77, 5.2). Epiphallus short and cylindrical 20 mm (19.2, 20.0). Flagellum short 36 mm (38, 32), nearly 1.2 times (0.68, 0. 87 ) the combined length of penis plus ‘epiphallus. Atrium abbreviated and ree ly distin- guishable. Lower vagina 5.82 mm (4.29, 5.32), exte nding to region of dart sacs; fa sub-equal dart sacs ds] (3.22 mm), ds2 (3.73 mm), ds3 (2.81 mm) and ds4 (2.30 mm) (2.39, 2.47, 1.98, 1.68) (2.35, 2.95, 2.81, 2.56). Dart bulbs embedded in vagina wall, forming conspicuous bulges. Median vagina sibbrreutetedl, bearing ring of four aia glands, dae glands reaching maximum height of 3.80 mm (3. 05-3.39) arn separatec alarm dart sacs =by distance of 1.83 mm length (2.86-2.81). Spermathecal duct short, 47mm (37-50) and without a caecum. Spermatheca elongated and sac-shaped, 9 mm length (7.54—9.81).
Type Material: Holotype: CNMO 7563; collected 9 August 2007 by Omar Mejia. Paratypes: CNMO 7564 (2): same data as the holotype. All from type locality.
O. Mejia et al., 2018
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10 mm
Figures 5-8. Humboldtiana aurea new species. Holotype, CNMO 7563. 5) Apertural view 6) Apical view 7) Oblique basal view 8)
Detail of embryonic whorls.
Type Locality: Chihuahua, E] Oro, municipality of Casas Grandes, near the border with the State of Sonora, 67 km southwest of Nuevo Casas Grandes, Chihuahua, 2054 m altitude (30°09’ 35” N, 108°33'56” W). The specimens were collected on the rocks next to a small waterfall, where the predominant vegetation is an oak-pine forest.
Distribution: Only known from the type locality.
Remarks: Humboldtiana aurea new species belongs to the H. texana species group based on its dart glands separated from the dart sacs. Two other species of the group had been described from the state of Chihuahua, H. balanites (Thompson, 2006) and H. corruga (Thompson and Mejia, 2006). Humboldtiana aurea is similar to H. balanites as both have a thin shell, an abbreviated atrium, and bulges on the dart sac caused by the underlying dart bulbs; nevertheless, the light shell color with dark-brown
bands of H. aurea contrasts with the dark-brown ground color with black bands of H. balanites. In addition, H. balanites has a multilobed verge, a long flagellum two times longer than the combined length of the penis plus epiphallus, a spermathecal duct that is twice the length of that present in H. aurea and, last but not least, H. balanites presents a short caecum in the spermathecal duct that is absent in H. aurea. On the other hand, H. aurea resembles H. corruga in size, but the shells of H. corruga are darker in color. The reproductive anatomies of H. corruga and H. aurea are similar in that both species have ware that protrude at the basis of the dart sacs, and both also have a short flagellum that is of same size as the combined length of the penis plus epiphallus; nevertheless, H. corruga differs from H. aurea by having a slenderer atrium, larger penis, dart sacs of the same size, a globular
spermatheca, and a spermathecal duct that with a caecum.
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THE NAUTILUS, Vol. 132, No. 3-4
Etymology: The specific epithet, aurea, alludes to the name of the type locality, “El Oro” (“The Gold”) where the species was collected.
Humboldtiana paquimei new species Mejia and Lépez (Figures 9-12, 15)
Diagnosis: Medium-sized Humboldtiana. The new species has a similar shell to H. regula (Thompson, 2006), but differs in reproductive anatomy particularly on the verge, the length of the epiphallus and the spermathecal ain. The description is based on the holotype and one paratype.
Description: SHELL (Figures 9-12) large, external lip not reflected, background color pale- Prous becoming darker toward aperture. Three well-defined chestnut- to dark- brown bands, third one wider than first and second. 4.1 Whorls (4.1). sEmbsyonte shell lighter than remainder of 2 (1.4) whorls and with sculpture of weak
shell, with 1
wavy striations. Sculpture of second whorl with small granules: remainder of shell with striations and white eerie Umbilicus almost completely covered by basal portion of ae Callus very thin, transparent. Shell height 37 mm (31); shell diameter 38 mm (33); aperture height 25 mm (21); aperture diameter 23 mm (20).
REPRODUCTIVE ANATOMY (Figure 15): Penis short 7.91 mm (7.24), slender at base and swollen in middle; short and stout verge of approximately one-third penis length and consisting of two tissue folds with digitiform processes. bens retractor muscle 3.04 mm long (7.7). Epiphalus long, cylindrical and stout, measuring 34 mm (29), four times penis length. Flagellum short 60 mm in length (62), nearly 1.4 times (1.7) eombined length of penis plus epiphallus. Atrium short and bare ly perceptible. Lower vagina of almost same length as penis, measuring 6.13 mm (5.68) and extending to region of dart sacs; four dart sacs of different sizes, ds] largest, remainder decreasing in size,
11
Figures 9-12. view 12) Detail of embryonic whorls.
V2
Humboldtiana paquimei new species. Holotype, CNMO 7565. 9) Apertural view 10) Apical view 11) Oblique basal
O. Mejia et al., 2018
Ds > IC Page 129
gen atr =| 5mm Figure 13. Reproductive anatomy of Humboldtiana wixarika
new species. Abbreviations. cae :spermathecal cae cum; dgla: dart glands;ds; dart sacs; epi: epiphallus; fla: flagellum; gen atr; genital atrium pen: penis; pr: penis retractor; spd: spermathecal duct: spt: spermatheca; vag: vagina; vd: vas deferens.
spt
spd
ee
vd
pen
gen atr
Figure 14. Reproductive anatomy of Humboldtiana aurea new species. Abbreviations. dgla: dart glands;ds; dart sacs; epi: epiphallus; fla: flagellum; gen atr; genital atrium pen: penis; pr: penis retractor; spd: spermathecal duct; spt: spermatheca; vag: vagina; vd: vas deferens.
spt
fla
aan . i epi ds"
vag gen atr
‘18
Figure 15. Reproductive anatomy of Humboldtiana paquimei new species. Abbreviations. cae :spe srmathecal caecum: dgla: dart glands;ds; dart sacs; epi: epiphallus; fla: flagellum; gen atr; genital atrium pen: penis; pr: penis retractor; spd: spermathecal duct; spt: spermatheca; vag: vagina; vd: vas deferens.
dsl (4.12 mm), ds2 (3.51 mm), ds3 (3.02 mm) and ds4 (3.77 mm) (4.25, 4.08, 3.0, 3.5). Median vagina bearing four dart glands that form ring just above Gime sacs arg] wal maximum height of (3.07mm) (3.02). Spermathecal duct long, measuring 112 mm (103), bearing short caecum of 8 mm length (6). Spermatheca sac-shaped, adhered to albumen gland, measuring 7 mm (10).
Type Material: Holotype: CNMO 7565; collected 9 August 2007 by Omar Mejia; Paratype: CN MO 7566 (1): same data as the holotype. All from type locality.
Type Locality: Chihuahua, bridge over Rio Piedras Verdes. Approximately 1 km Southeast Bh El] Willy, municipality of Casas Grandes, Chihuahua, 1829 m altitude (30°10'29" N, 108°18'40" W). Individuals were collected on the south wall of the canyon that surround the Piedras Verdes river, the predominant vegetation is an oak-pine forest.
Distribution: Only known from the type locality.
Remarks: The wavy sculpture of the embryonic whorl and the granular sculpture restricted to the first post- enilraworante assigns the new species to the subgenus Clydonac me, a taxon so far limited to the State of Chi- huahua that comprises six currently recognized species (Thompson, 2006). The shell of the new species is similar to H. regula (Thompson, 2006), but differs in three re- productive traits. In H. paquimei, the verge is stout and consisting of two tissue folds, in contrast to H. regula, where the verge is composed of a thin skirt of tissue. In
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THE NAUTILUS, Vol. 132, No. 3=4
H. paquimei, the epiphallus is four times larger than the penis, and in H. regula, the epiphallus is twice the length of the penis. Last but not least, in H. paquimei, the spermathecal duct plus the spermatheca is three times longer than in H. regula. Two shells deposited at the Florida Museum of Natural History (UF 185700 and UF103138) may be H. paquimei based on geographical proximity. These two specimens unfortunately alk reproductive anatomy and were excluded from the de scription.
Etymology: Paquimé was an ancient culture that de- veloped in northwestern Mexico between the years 700 and 1475. The maximum expression of this culture. is found in the municipality of Casas Grandes, Chihuahua. The name of the new species honors this culture and the archaeological zone of Paquimé, Intangible Cultural Heritage of Humanity since 1998.
ACKNOWLEDGMENTS
We are grateful to César Guzman for taking the photo-
graphs, to Alejandra Rocha- poe z for the illustrations of
the reproductive anatomy, to John Slapcinsky at Uni- versity of Florida Museum a Zoology who provided some photos of H. regula and two anonymous reviewers for their comments. This work was pé arti: ly funded by Conacyt project number 165990. This work is dedicated to the me mory of Prof. Fred G. Thompson.
LITERATURE CITED
Burch, J.B. and F.G. Thompson. 1957. Three new Mexican land snails of the genus Humboldtiana. Occasional Papers of the
Museum of Zoology, University of Michigan 590: 1-11: pls. ]—4.
Mejia, O. and G. Zuniga. 2007. Phylogeny of the three brown banded land snail genus Humboldtiana (Pulmonata: Humboldtianidae). Molecular Phylogenetics and Evolution 45: 587-595.
Pilsbry, H.A. 1927. The structure and affinities of Humboldtiana and related helicid genera of Mexico and Texas. Pro- ceedings of the Academy of Natural Sciences of Phila- delphia 79: 165-192.
Pilsbry, H.A. 1948. Inland Mollusca of Northern Mexico. I. The genera Humboldtiana, Sonorella, Oreohelix and Ashmu- nella. Proceedings of the Academy of Natural Sciences of Philadelphia 100: 185-203: pls. 12-14.
Thompson, F.G. 2006. A new land snail of the genus Humboldtiana (Gastropoda: Pulmonata: Humboldtia- nidae) from Nuevo Leén, Mexico. The Nautilus 120: 21-24.
Thompson, F.G. 2006. Some landsnails of the genus Hum- boldtiana from Chihuahua and Wester Texas. Bulletin of the Florida Museum of Natural History 46: 61-98.
Thompson, F.G. 2011. An annotated che cklist and bibliography of the land and freshwater snails of Mexico and Central America. Bulletin of the Florida Museum of Natural History 50: 1-299.
Thompson, F.G. and G.P. Brewer. 2000. Landsnails of the genus Humboldtiana from Northern Mexico (Gastropoda, Pulmonata, Helicoidea, Humboldtiani- dae). Bulletin of the Florida Museum of Natural History 43: 49-77.
Thompson, F.G. and O. Mejia. 2006. Two new land snails of the genus Humboldtiana (Gastropoda: Pulmonata: Humbold- tianidae) from Chihuahua, Mexico. The Nautilus 120: 95-99.
Von Ihering, H. 1892. Morphologie und Systematik des Geni- talaparates von Helix. Zeitschrift fiir Wissenschaftliche Zoologie 54: 425-520.
NAUTILUS
Volume 132
2018 AUTHOR INDEX
ANB BAU E at LD) Seetewe tee een OR ee ont oe aa tet ane ee Tee ae eran Ss neta 101 TER NAIN Eg ge retrer coments a Seth ee te SM Sars Src dav cesses aeteaavietaeeti- 51 /NIBGINUING), TRS -Secosenectbescnonedabssobucc coe ncceoDae eee Bee Cece ec eoer ne ees ROSE ane 19 EAGTETSSS) LITINUA, IMs “aoscecnconsanssesce soacde0 bodes SacBee A onecccl AE ooC AOA AEERACOOMESSRE 45 JAIVNINTOWR ISS oeconsnonceoroacedo testcase Bese nec enc aacane Recon c ee cece eee eeeenee 117 |EGISISZA® 18}.. coneaqposaanaescoosbocaceedoaadsoseLcesacen cits ape sess oSionneEe Bacco EOE 124 IBYOLGAINIE: Ns} eacdiadseodse dap tooncecaactoeticSentecibaSoeed ee ouceadaactodce nee eceaetnaees 45 NM Ne BUA eK O) otter cect onckdeciareceonee te acan eetnccr ocece at arese cio esse neers 124 IENORRIUATOWA, Ins -csesepsadticoadasockeesaae sondasshidececbodesaconsaouctenecuceeeese aarere 45 COOLING ACID Chat coed eaanccac cece eo poact ead tame sen OsaetE once Das REB CEE Pee 19 JBSIOTRINE, Ro, Seneasccsdanacaeeapncseonccesict ie aaaadicaodeaaneanoced Raaarasaaa eee meaner 33 IRUESWESS= (© @ NATE: FU easement ete ee 124 (GANSINAINO) (GANG [1S )o:. dchasaasstacatece eid oSecBee caEEe Bo ond eOCOS SOC CaS aaa REE 65 SAILILIBG. JNA CAN. Gk op sbansopaaescescos Boasecasscocuce ease Gack Coc eso Sa SeSCEECEEE 19 (CAIMIPANGIIATRT, 18%, accsondanodancsnsbocsecsadosuenactionnenc onuabdo nado OnE aT see EEE HEE §3 SUIMIOINTS: |] Lal RGILS cscoosdboaancteoccucebaecsosoaoer ase cBaenon ace ene ase onaeaaaceeee 101 CAWATITEATR Teg IB) Gara or essessacens Seas cons Ste seontveemeenrsdcsesscoyuesteaceeree 101] STARGINS Reh esses seortnss vis stare chasasareneemenes tet Soertantierovtotnre sain: | IDYOTUIGILAGS Sis. sosodenssandtoasesc5szeces seed neaeaee ss86oe ASSOUSSE EE SEES Pe anEcERRee 30 SONUZZIN, SAIS ia cacacectauc a ecniG i omaeene a shae ber oo eee O ne OOD Oc ERO 65 FAR © 1 ya Ege octet Bale wns cena nae ede ea ss the bare oa SoS 45 S OUMRES RS tiga serene tee ere eee te aban eRe rare er ol, 91 (ETOGI AR, ID) Uns Sacccocsacese eS Ese Tce e ce Oe SSN SOR ees E CoE ee 83 AMEN ORI OMIM Se ccccsescvsces esters ses sce seat sees senses sor nesevsses os sess seaesoes: 71 | SUATRACTERIAKOIRL, IMIG... conaonesseconscnosonoooncascoobeondesossonoebecooasebed SD, Uf UTE WAINNGY | F cieetcecnarstcgne voces tevesecyaecdudarsé cin? savodeeesn scadeaseecocieaes 30 USS ANDIDIAHICZZ ISG" acooceensoseacrosoobacecnadoenbedonask onc sonatanse opeconOs Oe Sea 117 AVES iad De syeabe stich So, Santen oder ci osha ki aaeeateneencmneaesieeahLcdeessieeeet: 08, 113 IJIN SORE (G8 Se oe erence oot coe Sete so loco, ucts Sadacecuacle st 117 LITAIN Ga | [eager ce eects Ase deetea Rare ese ous tages so. shore sen acest eben: 58 IKOATRYATITAWIE AIA pee tetra teeter ee rca ene Serre Rae cee er eet ccas Pastore teeta: 45 TABUNNNGE SNE DN aceettoccrces ecco ee eee eee Rc eee ae 13, 113 ILHANUVILIIG: (Gs Reds badconesosconespessonsue aac ROO SSNSODE coS NCEE EOS Deore 30 CSEVIAIN (Ca SU incest eee Oe oes DS ee Rite ila re a 13, 58 NEW TAXA PROPOSED IN VOLUME 132 GASTROPODA Boreocomitas inouei Amano, Hryniewicz, and Jenkins, 2018 (Pseudomelatomidae, fossil) .............00cccccceeeceeees settee eee eeeeeeerereeeeeeeees 119 G@alliostomassnesaaAnancmWeimandeAhnanes2 OlSosmewaspeciess Calliostomatidac) pes eee 62 Calliotiopissuapensise7nanveandeAnanees? 0ommewaspeciess (Calliotropid ac) pmseeeeeee eee eee 16 Humboldianaaurea Mejiarandeltonezs 2018 tmnewsspecies | (Elumboldtianid ac) esses sees esos eee ee ee 126 Humboldtiana paquimei Mejia and Lépez, 2018, new species (Humboldtianidae) ................c.cccececees ec eeeeeeseeeeeeeeeseseseseseeeesesenesenees 128 Humboldtiana wixarika Mejia, Lopez, and Reyes-Gémez, 2018, new species (Humboldtianidae)..........0..0.0.0:.0c cee eee 125 Memecilok-hanesupinzacw/nan teand Vel 0ommevvas pe cies (Gancel lari dae) pees eee nena tenn ee 114 Murdochella tricingulata Campagnari and Geiger, 2018, new species (Epitoniidae) ...........0..ccs cece eseeee ees ests esses esse eseseeeeesereseeeey 85 Reonidla Umanorum Sowines, AUS, messy GoSHES (ACBOMCS, HOSS) scccccosnce3e90005¢560580006003050000857505 65595345 0055650000 509093555050 OOCIGICEE 54 BIVALVIA
Cyclonaias necki Burlakova, Karatayev, Lopes-Lima, and Bogan, 2018 (new species) (Unionidae) ........0...0.00c0:cce cece eects 46
Beu, Alan
Bogan, Arthur E. Carew, James Cowie, Robert H. Del Rio, Claudia Julia deMaintenon, Marta Eernisse, Douglas Fedosov, Alexander Ford, David Garner, Jeff
Groves, Leslie
REVIEWERS FOR VOLUME I:
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