CRUSTACEA FROM THE MUHI QUARRY (ALBIAN–CENOMANIAN), AND A REVIEW OF APTIAN MECOCHIRIDAE...

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CRUSTACEA FROM THE MUHI QUARRY (ALBIAN–CENOMANIAN), AND A REVIEW OF APTIAN MECOCHIRIDAE (ASTACIDEA) FROM MEXICO RODNEY M. FELDMANN Research Associate, Section of Invertebrate Paleontology; Department of Geology, Kent State University, Kent, Ohio, 44242 ([email protected]) FRANCISCO J. VEGA Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Coyoacán, México, D.F. 04510, Mexico, ([email protected]) LETICIA MARTINEZ-LOPEZ Facultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico KATIA A. GONZÁLEZ-RODRÍGUEZ Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo, Km. 4.5 carretera Pachuca-Tulancingo, Pachuca, Hidalgo, 42184, Mexico OSCAR GONZÁLEZ-LEÓN Facultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico MARÍA DEL ROSARIO FERNÁNDEZ-BARAJAS Facultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico ABSTRACT The first record for the shrimp genus Aeger Münster, 1839, in America is documented by Aeger hidalguensis, new species, described from Albian–Cenomanian beds of the El Doctor Formation, Muhi Quarry, Hidalgo, Mexico. Reconsideration of Aegeridae Burkenroad, 1963, results in placement of Acanthochirana Strand, 1928, within the family. Aeger hidalguensis is associated with the palinurid lobster, Palinurus sp., which represents only the second fossil record for the genus in America. Specimens collected from the Aptian San Juan Raya Formation, Puebla, allow a more complete description of the mecochirid Meyeria pueblaensis Feldmann et al., 1995, and documentation of sexual dimor- phism in the species, the first report of dimorphism in the genus. KEY WORDS:Crustacea, Aegeridae, Palinuridae, Mecochiridae, Cretaceous, Puebla, Hidalgo, Mexico INTRODUCTION ANNALS OF CARNEGIE MUSEUM VOL. 76, NUMBER 3, PP . 145–156 31 OCTOBER 2007 In recent years, the number of species of fossil deca- pod crustaceans from Mexico has increased notably. This has prompted enthusiasm in several paleontolo- gists all around the country. Colleagues from other campuses of the Universidad Nacional Autónoma de México (UNAM), and from other Mexican universi- ties, have prospected new and well known localities and collected samples of undescribed forms, as well as of species previously reported. They have found specimens whose preservation quality and complete- ness permit a more detailed knowledge of their respective species. That is the case of two Cretaceous assemblages from Hidalgo and Puebla, central Mexico (Fig. 1). The Muhi Quarry is a recently discovered fossilif- erous locality within the Upper Cretaceous calcareous sequence of the Sierra Madre Oriental in Hidalgo (Fig. 2). It is located at 20° 45' 12.5" N and 99° 20' 29.4" W (Fig. 1). The fossils were collected from a light gray micritic limestone sequence with intercalat- ed black chert lenses that form part of the Albian–Cenomanian El Doctor Formation (Wilson et al. 1955; Segestrom 1956, 1961). Invertebrate remains include miliolids, ammonites, and comatulid and echinoid echinoderms, but the most conspicuous remains are diverse fish species. The presence of planktonic foraminiferans within the limestone, in association with the black chert lenses with radiolari- ans, suggests an open marine basin with a hypersaline and/or poorly oxygenated bottom (González- Rodríguez and Applegate 2000; González-Rodríguez et al. 2000, 2004, 2005; González-Rodríguez and Bravo-Cuevas 2005; González-Rodríguez and Fielitz 2005). Four specimens of Aeger hidalguensis, new species, and one specimen of Palinurus sp., are the crustacean representatives from the Muhi Quarry. San Juan Raya is a well-known locality among Mexican paleontologists. An Aptian age has been pro- posed and confirmed for the San Juan Raya Formation, whose geology and fossil content of

Transcript of CRUSTACEA FROM THE MUHI QUARRY (ALBIAN–CENOMANIAN), AND A REVIEW OF APTIAN MECOCHIRIDAE...

CRUSTACEA FROM THE MUHI QUARRY (ALBIAN–CENOMANIAN), AND A REVIEW OF APTIANMECOCHIRIDAE (ASTACIDEA) FROM MEXICO

RODNEY M. FELDMANNResearch Associate, Section of Invertebrate Paleontology;

Department of Geology, Kent State University, Kent, Ohio, 44242 ([email protected])

FRANCISCO J. VEGAInstituto de Geología, Universidad Nacional Autónoma de México,

Ciudad Universitaria, Coyoacán, México, D.F. 04510, Mexico, ([email protected])

LETICIA MARTINEZ-LOPEZFacultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico

KATIA A. GONZÁLEZ-RODRÍGUEZCentro de Investigaciones Biológicas, Universidad Autónoma del Estado de Hidalgo,

Km. 4.5 carretera Pachuca-Tulancingo, Pachuca, Hidalgo, 42184, Mexico

OSCAR GONZÁLEZ-LEÓNFacultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico

MARÍA DEL ROSARIO FERNÁNDEZ-BARAJASFacultad de Estudios Superiores Iztacala, UNAM, Tlalnepantla, Estado de México, 54070, Mexico

ABSTRACT

The first record for the shrimp genus Aeger Münster, 1839, in America is documented by Aeger hidalguensis, new species, described fromAlbian–Cenomanian beds of the El Doctor Formation, Muhi Quarry, Hidalgo, Mexico. Reconsideration of Aegeridae Burkenroad, 1963,results in placement of Acanthochirana Strand, 1928, within the family. Aeger hidalguensis is associated with the palinurid lobster, Palinurussp., which represents only the second fossil record for the genus in America. Specimens collected from the Aptian San Juan Raya Formation,Puebla, allow a more complete description of the mecochirid Meyeria pueblaensis Feldmann et al., 1995, and documentation of sexual dimor-phism in the species, the first report of dimorphism in the genus.

KEY WORDS:—Crustacea, Aegeridae, Palinuridae, Mecochiridae, Cretaceous, Puebla, Hidalgo, Mexico

INTRODUCTION

A N N A L S O F C A R N E G I E M U S E U M

VOL. 76, NUMBER 3, PP. 145–156 31 OCTOBER 2007

In recent years, the number of species of fossil deca-pod crustaceans from Mexico has increased notably.This has prompted enthusiasm in several paleontolo-gists all around the country. Colleagues from othercampuses of the Universidad Nacional Autónoma deMéxico (UNAM), and from other Mexican universi-ties, have prospected new and well known localitiesand collected samples of undescribed forms, as wellas of species previously reported. They have foundspecimens whose preservation quality and complete-ness permit a more detailed knowledge of theirrespective species. That is the case of two Cretaceousassemblages from Hidalgo and Puebla, centralMexico (Fig. 1).

The Muhi Quarry is a recently discovered fossilif-erous locality within the Upper Cretaceous calcareoussequence of the Sierra Madre Oriental in Hidalgo(Fig. 2). It is located at 20° 45' 12.5" N and 99° 20'29.4" W (Fig. 1). The fossils were collected from alight gray micritic limestone sequence with intercalat-

ed black chert lenses that form part of theAlbian–Cenomanian El Doctor Formation (Wilson etal. 1955; Segestrom 1956, 1961). Invertebrateremains include miliolids, ammonites, and comatulidand echinoid echinoderms, but the most conspicuousremains are diverse fish species. The presence ofplanktonic foraminiferans within the limestone, inassociation with the black chert lenses with radiolari-ans, suggests an open marine basin with a hypersalineand/or poorly oxygenated bottom (González-Rodríguez and Applegate 2000; González-Rodríguezet al. 2000, 2004, 2005; González-Rodríguez andBravo-Cuevas 2005; González-Rodríguez and Fielitz2005). Four specimens of Aeger hidalguensis, newspecies, and one specimen of Palinurus sp., are thecrustacean representatives from the Muhi Quarry.

San Juan Raya is a well-known locality amongMexican paleontologists. An Aptian age has been pro-posed and confirmed for the San Juan RayaFormation, whose geology and fossil content of

corals, gastropods, ammonites, bivalves, brachiopods,echinoderms, and dinosaur footprints have been stud-ied by several authors (Nyst and Galeotti 1840;Villada 1905; Aguilera 1906; Burckhardt 1930;Müllerried 1934; Lambert 1935; Alencáster 1956;Calderón 1956; Reyeros 1963; De Cserna 1969, 1970;Buitrón 1970; Ferrusquía 1970; Buitrón and Barceló1980; Rodríguez de la Rosa et al. 2004; among oth-ers). Feldmann et al. (1995) described Meyeriapueblaensis based on four specimens collected fromtwo localities of the San Juan Raya Formation. Newmaterial collected from one locality at 18° 19' 29.9" Nand 97° 54' 20.6" W (Fig. 3) is represented by sixnearly complete specimens and four fragmentaryspecimens, preserved within a gray mudstone.Additionally, two new specimens of the species werecollected from locality B of Feldmann et al. (1995).Paleoenvironment for these deposits has been inter-preted by them as a shallow lagoon.

Specimens are deposited in the Colección delLaboratorio de Geología-Paleontología, Facultad deEstudios Superiores Iztacala, Universidad NacionalAutónoma de México, D. F., under the acronym IZT-M;in the Museo de Paleontología, Centro deInvestigaciones Biológicas, Universidad Autónoma delEstado de Hidalgo, under the acronym UAHMP; and inthe Section of Invertebrate Paleontology, CarnegieMuseum of Natural History, Pittsburgh, Pennsylvania,U.S.A., under the acronym CM.

SYSTEMATIC PALEONTOLOGY

Order Decapoda Latreille, 1802Suborder Dendrobranchiata Bate, 1888

Superfamily Penaeoidea Rafinesque, 1815Famly Aegeridae Burkenroad, 1963

Included genera.—Aeger Münster, 1839; Acanthochirana Strand,1928.

Genus Aeger Münster, 1839

Type species.—Macrourites tipularius Schlotheim, 1822, by subse-quent designation of Woods (1925).

Aeger hidalguensis, new species(Fig. 4a–c; Fig. 5a–f)

Types.—Holotype, UAHMP-2246, paratypes, UAHMP-615,UAHMP-711, UAHMP-824.

Diagnosis.—Medium sized aegerid, cephalothorax subtriangular,abdomen longer than cephalothorax. Third maxilliped more than twicetotal carapace length, ornamented with long and thin spines on loweredge.

Description.—Medium sized aegerid, cephalothorax subtriangular;

hepatic groove short; branchial chambers ovoid-elongate. Abdomenshorter than or equal to cephalothorax length; abdominal somites 1 and2 shortest, abdominal somite 6 longest; pleura subtriangular elongatedin somites 2–5; pleopod basis appears to be subtriangular. Telson 0.33abdomen length, exopodites and endopodites of telson suboval, withmedian groove. Epistome subtriangular, with two sharp teeth.Antennae twice total body length. Third maxilliped long, 2.7cephalothorax length; double row of spines on lower side of articles.Ischium subtrapezoidal; merus semirectangular, elongate, five times aslong as ischium; carpus subrectangular, half the length of merus, lowersurface convex, rows of seven spines on each side; propodus verylarge, subrectangular, two and one half times as long as carpus, lowersurface concave, with two rows of spines; dactylus short and thin, withat least three spines on lower surface and two on apex. First pair ofpereiopods chelate; other pereiopods long and thin.

Measurements.—UAHMP-2246: cephalothorax length = 27.5 mm,height = 11.0 mm; abdomen length = 13.6 mm, height = 7.7 mm; 3rdmaxilliped length = 77.8 mm, height = 3.1 mm; antennae length = 33.4mm. UAHMP-615: cephalothorax length = 28.6 mm, height = 12.8mm; abdomen length = 26.8 mm, height = 11.5 mm; antennae length =80.5 mm. UAHMP-711: cephalothorax length = 31.9 mm, height =20.3 mm; abdomen length = 32.0 mm, height = 17.5 mm; 3rd maxil-liped length = 50.8 mm. UAHMP-824: cephalothorax length = 26.2mm, height = 12.8 mm; abdomen length = 21.4 mm, height = 11.5 mm;antennae length = 70.7 mm.

Etymology.—The trivial name refers to the state ofHidalgo, from which the specimens were collected.

Occurrence.—Muhi Quarry, at 20° 45' 12.5" N and 99° 20' 29.4" W,Hidalgo, Mexico; El Doctor Formation, Albian–Cenomanian.

Discussion.—The Aegeridae has typically been thoughtto include only Aeger; however, if it is defined asembracing genera characterized by long third maxil-lipeds bearing movable spines, the first three pereiopodslong, slender, and chelate, and accessory spines on atleast some of the pereiopods, then AcanthochiranaStrand, 1928, must also be placed within the family. Byimplication, that is the position taken by Etter (2004)when he compared details of morphology of Aeger andAcanthochirana. Other authors have placed the generawithin the Penaeidae (Garassino and Teruzzi 1990;Garassino 1994). The problem of making clear distinc-tions in family placement is caused in large part byincomplete specimens and poor preservation of the very

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Fig. 1.—Location map of study areas in Mexico.

delicate cuticle comprising the exoskeleton. The speci-mens from Hidalgo are no exception with regard topreservation. The carapace and abdomen are severelycrushed and the details of the groove pattern and orna-mentation on the cephalothorax are almost totallyobscured; however, the third maxilliped is well pre-served and there can be no doubt that the specimens arereferrable to the Aegeridae and the genus Aeger.

Species of Aeger can be distinguished from species ofAcanthochirana in several ways, clearly summarized byFörster (1967) (Table 1). With regard to A. hidalguensis,the relative lengths of abdominal somites 1/2 and 5/6are consistent with placement in Aeger (Table 1).Although not noted in the table, the third maxilliped isalso much shorter in Acanthochirana than in Aeger.Although Förster (1967, p. 164) and Schweigert (per-sonal communication) cautioned that small individualstend to have proportionately longer third maxillipeds,the extreme length of that appendage in A. hidalguensisstrongly suggests placement of the species in Aeger.Acanthochirana smithwoodwardi (Van Straelen, 1940)has a third maxilliped as long as the carapace(Garassino 1994), for example. Principal differencesbetween Acanthochirana and Aeger are discussed inEtter (2004), who also commented on Aeger sp. fromthe Jurassic of northern Switzerland and compared it tomost other species within the genus. None of thespecies Etter considered has a third maxilliped as longas that exhibited by A. hidalguensis. Comparisons with

Aeger tipularius (Schlotheim, 1822), the type species,discussed by Schweigert (2001), are difficult to discernbecause of the preservational quality of A. hidalguensis,but the carapace of the latter species seems to be pro-portionately more elongate and the length of third max-illiped is much longer in the Mexican specimens than inthe type species. The specimen illustrated by Barthel etal. (1990, p. 133) is not the type species as, indicated,but rather should be referred to Aeger spinipes(Desmarest, 1822) (Schweigert 2001). The onlyCretaceous species is Aeger libanensis Roger, 1946,from the Cenomanian of Lebanon, which has very longpereiopods, but the third maxilliped is not as long as

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Fig. 2.—Geological map of the vicinity of the Muhi Quarry, Hidalgo. Map by Víctor Bravo-Cuevas.

Fig. 3.—Location map of locality in San Juan Raya, Puebla.

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Fig. 4.—a–c, Aeger hidalguensis, new species. a, reconstruction based on holotype UAHMP-2246; b, reconstruction of anterior part of cephalothorax,based on holotype UAHMP-2246; c, reconstruction of cephalothorax, based on paratype UAHMP-824; d–h, Meyeria pueblaensis Feldmann, Vega,García-Barrera, Rico-Montiel and Martínez-López, 1995; d, reconstruction based on hypotype IZT-M-3A; e, abdomen based on hypotype IZT-M-10A;f, right first pereiopod, based on hypotype IZT-M-11; g, reconstruction of telson, based on hypotype IZT-M-17; h, reconstruction based onhypotype;IZT-M-9; i, Palinurus sp. Reconstruction based on hypotype UAHMP- 659. Abbreviations for morphological features: Ab = abdomen; An =antennae; BrG = branchial groove; CG = cervical groove; Cph = cephalothorax; Ep? = epistome?; Epth? = epistome teeth?; HG = hepatic groove;Max3 = 3rd maxilliped; Mpt? = mouth parts; Pe1 = first pereiopod; Pl = pleura; Te = telson; Ur = uropod.

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Fig. 5.—a–g, Aeger hidalguensis, new species. Muhi Quarry, Hidalgo. Albian–Cenomanian. El Doctor Formation. a, specimen preserving left thirdmaxilliped (note spines on ventral surface), holotype UAHMP-2246; b, specimen preserving large antennae [Note possible fragment of third maxil-liped and comatulid arm.], paratype UAHMP-824; c, specimen preserving long antennae and some pereiopods, paratype UAHMP-824; d, nearly com-plete specimen [Note long third maxilliped.], paratype UAHMP-711; e, detail of spines on ventral surface of right third maxilliped, paratype UAHMP-711; f, detail of abdomen (note pleura), paratype UAHMP-711; g, Palinurus sp., hypotype, UAHMP-659. Muhi Quarry, Hidalgo. Albian–Cenomanian,El Doctor Formation. Scale bars equal 10.0 mm.

that in A. hidalguensis. Thus, the differences notedbetween previously described species of Aeger and theMexican specimens, including the length of thepereiopods and size of the third maxilliped, warrant thenew species.

Although species of Aeger have been noted previ-ously from Late Cretaceous occurrences, none has beenrecorded from the Western Hemisphere. Thus, theoccurrence in Mexico represents a significant expansionof the biogeographic range of the taxon.

Suborder Pleocyemata Burkenroad, 1963Infraorder Astacidea Latreille, 1802

Family Mecochiridae Van Straelen, 1925

Genus Meyeria M'Coy, 1849

Type species.—Astacus ornatus Phillips, 1829, by original designa-tion.

Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico-Montiel and Martínez-López,

1995(Fig. 4d–h; Fig. 6a–g; Fig. 7a–g, Fig. 8a–c)

Types.—Hypotypes, CM 55269-55275, IZT-M-2A, IZT-M-3A, IZT-M-3B, IZT-M-4, IZT-M-7, IZT-M-9, IZT-M-10A, IZT-M-10B, IZT-M-11, IZT-M-12, IZT-M-16, IZT-M-17, IZT-M-23.

Emended diagnosis.—Medium sized mecochirid, males much largerthan females; cephalothorax shorter than abdomen. Cervical groovedeep, hepatic groove subparallel to anteroventral margin. Rostrum tri-angular. Abdomen surface smooth; pleura of male abdominal somites

subrectangular with posteroventral spine, ornamented with fine spineson their edges; pleura of female abdominal somites ovoid. Uropodssubquadrate, with diaeresis; region of uropods beyond diaeresis veryfinely ribbed longitudinally, possibly uncalcified. Lower surface ofpereiopods covered by double row of granules.

Description.—Medium sized mecochirid. Cephalothorax shorter thanabdomen. Cervical groove deep, inclined toward anteroventral margin.Hepatic groove subparallel to anteroventral margin, sinuous in junc-tion with straight branchiocardiac groove, inclined about 30° towardposterior margin. Rostrum straight, triangular

Abdomen 1.25 times length of cephalothorax, abdominal somitessmooth. Pleura of somites 2–5 ornamented with fine spines on theedge. Pleura of male somite 2 subrectangular, anterior margin round-ed, posterior margin straight; pleura of male somites 3 and 4 sub-quadrate; pleura of male somites 5 and 6 subrectangular, with roundedventral margins. Pleura of female somites 1–5 semioval, pleura offemale somite 6 subcircular. Uropods subquadrate, with diaeresis, lat-eral margins with four sharp spines; extension of uropods beyonddiaeresis and termination of telson with fine, longitudinal ribs, possi-bly weakly calcified.

Pereiopods slender, long, with tuberculate ventral margins. Firstpereiopods large, length 1.33 times length of cephalothorax; ischiumsubquadrate, narrows at junction with subrectangular merus; carpusnot clear; propodus elongate, rectangular, two sharp small spines nearjunction with uncinate dactylus. Pereiopods 2 and 3 of similar length,0.33 times shorter than first pereiopod; pereiopods 4 and 5 smaller,0.33 times shorter than pereiopods 2 and 3.

Measurements.—Measurements are presented for all specimens thatare well enough preserved to obtain significant values. Gender of thespecimen is indicated where it was possible to determine.

CM 55273: cephalothorax length = 9.4 mm, height = 7.5 mm;abdomen length = 14.3 mm, height = 3.5 mm.

CM 55269: male, cephalothorax length = 19.3 mm, height = 8.1mm; abdomen length = 27.0 mm, height = 12.4 mm; left first pereio-pod length = 26.9 mm.

CM 55271: female, cephalothorax length = 4.6 mm, height = 3.8mm; abdomen length = 11.7 mm, height = 3.1 mm.

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TABLE 1. Morphological features of Aeger and Acanthochirana that distinguish species of the two genera¹.

Aeger Acanthochirana

Dorsum of rostrum smooth Dorsum of rostrum dentateVenter of rostrum with infrarostral spine Venter of rostrum without infrarostral spinePostorbital spine behind orbital notch Postorbital spine absentAbdominal somite 1 short (S1/S2 < 0.7) Abdominal somite 1 only slightly shorter than S2 (S1/S2 > 0.8)

Abdominal somite 6 scarcely lengthened (S5/S6 = 0.9) Abdominal somite 6 strongly elongate (S5/S6 = 0.6 – 0.7)

In both genera, there is a relative size decrease of S6 with increase in size of individualsCarapace surface granular Carapace surface smoothAntennular flagellum longer than individual, Antennular flagellum short, generally shorter than cephalothorax

in young individuals up to 3 times carapace lengthMxp3, P1, and P2 with movable spines Mxp3 and only P1 with movable spinesCarpi of P2 and P3 moderately elongated Carpi of P2 and P3 strongly elongated

___¹(translated from Förster 1967, p. 168)Characters denoted in bold face type are evident on Aeger hidalguensis.S = abdominal somite; Mxp3 = third maxilliped; P = pereiopod.

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Fig. 6.—a–g, Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico-Montiel and Martínez-López, 1995. San Juan Raya, Puebla. Aptian, San JuanRaya Formation. a, nearly complete male specimen, hypotype IZT-M-3A; b, detail of cephalothorax, hypotype IZT-M-16; c, specimen preserving bothfirst pereiopods, hypotype IZT-M-4; d, specimen preserving several anterior pereiopods, hypotype IZT-M-12; e, detail of right first pereiopod, hypo-type IZT-M-11; f, detail of male abdomen [Note ornament on edge of pleura.], hypotype IZT-M-10A; g, specimen preserving telson, hypotype IZT-M-17. Scale bars equal 5.0 mm.

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Fig. 7.—a–g, Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico-Montiel and Martínez-López, 1995. San Juan Raya, Puebla. Aptian, San JuanRaya Formation. a, nearly complete specimen with first pereiopods, hypotype CM 55269; b, male specimen with nearly complete abdomen and par-tial cephalothorax, hypotype IZT-M-2A; c, partial cephalothorax and first pereiopods, hypotypes IZT-M-7; d, possibly decayed specimen, hypotypeIZT-M-23; e, specimen preserving abdomen (note semioval pleura) and right first pereiopod, hypotype IZT-M-9; f, specimen preserving abdomen (notesemioval pleura) and telson, hypotype CM 55271; g, nearly complete female specimen with left first pereiopod. Hypotype CM 55273. Scale bars equal5.0 mm.

IZT-M-2AL: male, cephalothorax length = 14.4 mm, height = 5.7mm; abdomen length = 27.8 mm, height = 9.3 mm.

IZT-M-3A: male, cephalothorax length = 23.0 mm, height = 13.1mm; abdomen length = 30.3 mm, height = 10.2 mm.

IZT-M-4: cephalothorax length = 18.7 mm, height = 7.8 mm;abdomen length = 25.5 mm, height = 7.0 mm; left first pereiopodlength = 27.3 mm.

IZT-M-7: right first pereiopod length = 28.7 mm. IZT-M-9: female, cephalothorax length = 6.0 mm, height = 5.6

mm; abdomen length = 12.8 mm, height = 3.9 mm. IZT-M-10A: male, abdomen length = 18.6 mm, height = 8.8 mm. IZT-M-11: first pereiopod length = 22.9 mm. IZT-M-12: cephalothorax length = 17.8 mm, height = 6.9 mm;

abdomen length = 23.1 mm, height = 5.1 mm. IZT-M-16: cephalothorax length = 11.5 mm, height = 7.5 mm. IZT-M-17: cephalothorax length = 21.0 mm; abdomen length =

15.9 mm; telson length = 7.5 mm. IZT-M-23: cephalothorax length = 21.8 mm, height = 16.4 mm;

abdomen length = 18.1 mm, height = 11.5 mm; first pereiopod length= 20.3 mm.

Occurrence.—San Juan Raya, at 18° 19' 29.9" N and 97° 54' 20.6" W,Puebla, Mexico; San Juan Raya Formation, Aptian.

Discussion.—The original description of Meyeriapueblaensis was based on the morphology of two smallspecimens. Representatives here described span a widerrange of sizes, although all tend to be smaller than otherspecimens assigned to species of Meyeria. The newmaterial also provides morphological details of the pleu-ra, telson, and pereiopods not available previously toFeldmann et al. (1995).

Perhaps the most significant observation is that theabdominal pleura exhibit two quite different forms. Thepleura on the larger specimens are generally quadrate andbear small spines on the posteroventral corner whereas

those on the smaller specimens are more rounded and lackthe posteroventral spines. This relationship of pleural mor-phology has been recognized previously (Étallon 1859;Forest and de Saint Laurent 1989; Feldmann and de SaintLaurent 2002) in living and fossil glypheids. In the livingform, Neoglyphea inopinata Forest, de Saint Laurent, andChace, 1976, the pleura of males were quadrate and spinedwhereas those of the females were more rounded. Étallon(1859) had drawn the same conclusion by studyingGlyphea regleyana (Desmarest, 1822), and Feldmann andde Saint Laurent applied these observations to deduce thegender of a specimen of Glyphea foresti Feldmann and deSaint Laurent, 2002. It seems reasonable to apply the samereasoning to specimens of M. pueblaensis. The speciesbecomes the first representative of Meyeria in which sex-ual dimorphism has been documented.

The four specimens judged to be males exhibit cara-pace size and shape differences from those of females.Carapace lengths of males are about twice that of the twospecimens confirmed to be females. Herrick (1911)observed that male Homarus americanus H. MilneEdwards, 1837, tend to be larger than females, but they arenot twice as large. In addition, the carapaces of males aremuch more slender that those of confirmed females.Unfortunately, the small sample of specimens of M.pueblaensis for which gender can be determined does notpermit distinguishing among true size differential, partialsampling of the total size ranges of males and females,sampling of different age classes, and length/height distor-tion due to compression. However, the very large differen-tials in dimensions make it clear that females of the speciesare smaller and stouter than males.

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Fig. 8.—a–c, Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico-Montiel and Martínez-López, 1995. San Juan Raya, Puebla. Aptian, San JuanRaya Formation. a, hypotype CM 55269, showing male abdomen indicated by acute terminations on abdominal pleura (arrows); b, hypotype CM55270, showing female abdomen indicated by rounded terminations on abdominal pleura (arrows) [Note the difference in scales of male and femalespecimens.]; c, hypotype CM 55271 showing longitudinally ribbed terminations on uropods (arrows). Scale bars equal 1 cm.

Infraorder Palinura Latreille, 1802Superfamily Palinuroidea Latreille, 1802

Family Palinuridae Latreille, 1802

Genus Palinurus Weber, 1795

Type species.—Astacus elephas Fabricius, 1787, by monotypy.

Palinurus sp.(Fig. 4i; Fig. 5g)

Type.—Hypotype UAHMP-659.

Description.—Cephalothorax subrectangular, covered by forward-directed sharp spines, uniformly spaced but more numerous behindcervical groove. Spine distribution in front of cervical groove formingan inverted U-shape; a perpendicular row extends from rostrum to thebase of spines oriented in U-shape. Cervical groove deep, interceptslateral margin in anterior one third of cephalothorax and crosses mid-line at about midlength of cephalothorax. Frontal horns triangular, pro-jected beyond anterior margin. Rostrum triangular, half the length ofrostral spines. Antennae covered with spines; left antenna length atleast 1.25 times cephalothorax length. Antennular peduncle subcylin-drical, 0.25 times maximum cephalothorax length.

Measurements.—Cephalothorax length = 35.0 mm, width = 26.8 mm;right antenna length = 37.1 mm, width = 7.2 mm.

Occurrence.—Muhi Quarry, at 20° 45' 12.5" N and 99° 20' 29.4" W,Hidalgo, Mexico; El Doctor Formation, Albian–Cenomanian.

Discussion.—The sole specimen probably represents amolt. The cephalothorax and abdomen are still articulat-ed, but no traces of appendages are seen. Palinuruspalaciosi Vega et al., 2006 from the Albian of Chiapas(Vega et al. 2006) is the first fossil record for the genusin America. It differs from the Hidalgo specimen in hav-ing weaker cephalothoracic spines, but stronger andlonger frontal horns. Other Palinurus fossil representa-tives are known from the Cretaceous (Turonian,Senonian) of Europe, from the Upper Cretaceous(Cenomanian) of Lebanon, and from the Tertiary (mid-dle Eocene) of Europe. Palinurus sp. from Hidalgo issimilar to Palinurus woodwardi Fritsch, 1887 (Turonianof Czech Republic), Palinurus baumbergicus Schlüter,1872 (Upper Cretaceous of Germany), and Palinurus sp.(Cenomanian of Lebanon, in Garassino 1994), in sizeand shape of the frontal horns, as well as in cephalotho-rax ornament. Other Lower Cretaceous Palinurus arefound in the upper Barremian–Aptian and Albian beds ofNE Italy (Bravi and Garassino 1998; Garassino 2000).The state of preservation in the Hidalgo specimen is notappropriate to make a detailed comparison with otherfossil species.

ACKNOWLEDGMENTS

Special thanks to Hiroaki Karasawa, Mizunami Fossil Museum,Mizunami, Japan, for his usual kindness and help. Advice of GünterSchweigert, Staatliches Museum für Naturkunde, Stuttgart, Germany,is appreciated. Comments regarding the placement of Acanthochiranaby Alessandro Garassino, Museo Civico di Storia Naturale di Milano,

and on aspects of sexual dimorphism by Frederick Schram, Langley,Washington, U.S.A., were extremely useful. Observations on genderdifferences in size of lobsters were discussed with J. S. Cobb,Kingston, Rhode Island, U.S.A. An early draft of the manuscript wasread and improved by Carrie Schweitzer, Kent State University, StarkCampus, North Canton, Ohio. Careful reviews by Dale Tshudy,Edinboro University of Pennsylvania, U.S.A., Guenter Schweigert,and an anonymous reviewer greatly improved the final work. Ourthanks to these individuals.

IN MEMORIAM

To the memory of our dear friends, colleagues and teachers, SheltonP. Applegate and Roberto Rico Montiel, who shared with us the enthu-siasm and passion for the study of fossils.

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