Contributions to Zoology, 67 (4) 257-266 (1998)

SPB Academic Publishing bv, Amsterdam

Late stomatopods (Crustacea, ) from Israel and

Jordan

Cees+H.J. Hof

Institute for Systematics and Population Biology, University of Amsterdam, P.O. Box 94766,

1090 GT Amsterdam, The Netherlands

Keywords: Stomatopoda, Mesozoic, Cretaceous, fossils, taphonomy, , Squilloidea, new

family Ursquillidae, new Ursquilla

Abstract fossils from Jordan and Israel, confirms Glaessner’s

suggestion. The fossils, originating from three dif-

The eryonid decapod Eryon yehoachi Remy & Avnimelech, allow ferent localities (Fig. 1), a detailed recon- the is 1955, from Late Cretaceous of Israel, redescribed as a struction of the telson, the sixth abdominal tergite, fossil stomatopod species within the new genus Ursquilla. This and parts of the uropods. Although this analysis is redescription is based on the original type specimen and two

based on fossil the telson of additional records from Israel and Jordan. The material allows incomplete material,

a detailed reconstruction of the telson, the sixth abdominal this Cretaceous stomatopod carries enough informa-

tergite, and part of the uropods. The distinct telson ornamen- tion to recognize a new genus within a new family. tation ofthese the erection of stomatopods justifies a new family The newly recognized stomatopod sheds a fresh within the superfamily Squilloidea. light on the evolution of these remarkable crusta-

ceans.

Introduction

Abbreviations of depositories In 1955, Remy & Avnimelech described Eryon

a new of the extinct yehoachi, species decapod BMNH, The Natural History Museum, London

GSI, family . Their description was based on Geological Survey of Israel, Jerusalem

MNHN, Museum National d’Histoire Naturelle, Paris a single specimen from the Upper Cretaceous (Cam-

panian) of the Ouadi Seiyal Valley in eastern Israel.

The authors interpreted the fossil remains as the Systematics of tergal plates an eryonid céphalothorax showing

the inner side. Although they comprehensively Class Malacostraca Latreille, 1806 justified the identification of the eryonid based only Subclass Hoplocarida Caiman, 1904 on the inner side of the céphalothorax, they seemed Order Stomatopoda Latreille, 1817 to have no doubt about the taxonomic accuracy of Suborder Unipeltata Latreille, 1825 their identification. Superfamily Squilloidea Latreille, 1803 However, Glaessner (1969: 470) suggested

that the E. description of yehoachi was actually based on a Squilla- like stomatopod telson. If Ursquillidae Hof, new family

Glaessner is right about the stomatopod affinities of this the material could fossil, provide impor- Diagnosis. - Moderate sized stomatopods. Telson

tant information the considering (late) Mesozoic about as long as broad with paired submedian-, radiation of the stomatopod taxa. intermediate-, and lateral teeth, and distinct lateral

A of the of E. folds re-study type specimen yehoachi at one third from the anterior margin. Four together with three similar rounded previously undescribed, intermediate marginal denticles present

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Ursquilla Hof, new genus

Diagnosis. - As for family.

Type species. - Eryon yehoachi Remy & Avni-

melech, 1955, by present designation and mono-

typy. Originally described as the cephalothorax of

an eryonid decapod.

- The Etymology. name is derived from the prefix

ur, meaning primeval, in combination with the

generic name Squilla.

Ursquilla yehoachi (Remy & Avnimelech, 1955)

new combination

(Figs. 2-4)

Synonymy, - Eryon yehoachiRemy & Avnimelech, 1955:311,

pi. 14a; Glaessner, 1969: 470 [squillid stomatopod telson, not

Holthuis & 1969: 541 Eryon]; Manning, [may belong to genus

Squilla ]; Yun, 1985: 19 [referring to Holthuis, 1969; should

be Holthuis & Manning, 1969],

Material examined. - The holotype ofEryon yehoachi (MNHN,

R, This shows the 62691) (Fig. 2A). specimen imprint of a

telson and part of the sixth abdominal tergite on a small slab

of yellowish chert. An additional specimen came from the

Geological of Israel (GSI, It con- I. for Survey M-8113) (Fig, 2C). Fig. Locality map Ursquilla yehoachi (Remy & Avnime- find cerns a relatively recent (1994) by Dr. S. llani. The spec- lech, 1955): 1) Israel, in the vicinity ofthe city ofArad (MNHN, imen shows the imprint of a telson, part ofthe sixth abdominal R. 62691), 2) Israel, north ofthe erosion cirque “Hamakhtesh tergite and fragments of both the uropods on a small piece of Haquatan”(GSI, M-8113), 3) Jordan, in the vicinity ofthe city chert. brown Two specimens on one slab originate from The of Karak (BMNH, I 7316). [Disputable state boundaries are Natural History Museum in London (BMNH, I 7316) (Fig, 3A). not indicated on this map.] Both these specimens show the incomplete imprint ofa telson,

one including the remains ofthe sixth abdominal tergite. The

telsons two are preserved at different levels on a slab of fine

grained, yellowish shaly limestone. The label relates that these and a linear row of altogether four rounded sub- specimens were presented by Dr. F. Johnson through Frank T. median denticles. Telson dorsally ornamented Ellis in 1906, In the collection of the BMNH May an epoxy with median carina and a paired intermediate-, resin mould ofthe from Paris type specimen is present, indexed lateral-, and marginal carinae. Median carina under number I 15472.

flanked by a string of tubercles. At the level of

Localities and lithostratigraphy. - Along with the description the telson’s lateral folds a low transverse carina of Etyon yehoachi, Remy & Avnimelech (1955) provide some connects the lateral carina with the marginal carina. information on the locality and the stratigraphical position of Between the lateral fold and the intermediate this fossil. The specimen was found in the southern part ofthe teeth the carinae marginal marginal carry minute, Judean Desert in the Ouadi Seiyal Valley (presently named

sharp, lateral tubercles. All carinae as well as the Nahal Ze’elim) along the route from Be’er She’va to Mount

Sedom. The specimen labelprovides a coarse locality indication submedian marginal teeth are rounded and orna- by referring to the coordinates 17./08. of the Israeli grid. mented with a dorsal row of tubercles. According to Remy & Avnimelech the specimen comes from

one ofthe Campanian chert banks of the alternating calcareous

- The new Type genus. monotypic genus Ursquilla, and siliceous beds covering the southeastern flanks of the

as defined below. anticline of the Judean mountains.

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Fig. 2. & fossil Ursquilla yehoachi (Remy Avnimelech, 1955) imprints from Israel with corresponding ammoniumchloride covered peels. A, Flolotype R. telson and sixth abdominal MNHN, 62691, showing somite; B, peel ofholotype; C, GSI, M-8113, showing sixth abdominal and of the ammonium telson, somite, parts uropods; D, chloride covered peel ofthe GSI specimen. Arrows indicate

of lateral Abbreviations ofcarinae of last abdominal = folds. = = position somite: sm submedian carinae, im intermediate carina, 1

= lateral carina, m carina. of = marginal Abbreviations uropodal structures: = = up uropodal protopod, ue uropodal exopod, un

outer of Abbreviations = = = possible margin uropodal endopod. for telson ornament: me median carina, bn basal notch. It line of

the median = = tubercles flanking carina, sc submedian Ic lateral mac = = carina, carina, marginal carina, smt submedian marginal

it = teeth, intermediate tooth, lat = lateral = shells. marginal marginal tooth, g gastropod Scale bars all 1 cm.

to Dr. Zeev ofthe of According Lewy Geological Survey Israel (lower ofthe Mishash which member) Formation, ranges from the (pers. comm.) fossil was most collected in the the likely vicinity uppermost Lower Campanian into the Middle Campanian. of the of Arad The to city (Fig. 1, locality 1). chert banks Biostratigraphically these banks are within the ammonite which Remy & Avnimelechrefer belong to the Chert Member Hoplitoplacenticeras marroti Zone (Reiss et al., 1986).

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Fig. 3. Two Ursquilla yehoachi (Remy & Avnimelech, 1955) fossil imprints from Jordan with corresponding ammonium chloride

and the half of telson at covered peels. A, BMNH, I 7316, showing a telson with attached sixth abdominal somite posterior a a

slightly higher level; B, ammonium chloride covered peel ofthe telson with attached sixth abdominal somite; C, ammoniumchloride

somite; = covered peel of the half telson. Arrows indicate position of lateral folds. Abbreviation of carinae of last abdominal sm

= = submedian carinae. Abbreviations for telson ornament: bn = basal notch median carina, sc submedian carina, lat lateral marginal

= = bars all 1 tooth, it = intermediate marginal tooth, imd intermediatemarginal denticles, smd submedian marginal denticles. Scale

cm.

of found east of Karak in of the Amman Silicified The GS1 specimen originates from the same Chert Member outcrops

found the Limestone Formation Zeev the Mishash Formation; only this specimen was at (pers. comm. Lewy).

northern flank of the Hazera Anticline, north of the erosion

- Three cirque “Hamakhtesh Haquatan” (Fig. 1, locality 2). The speci- Description. of the four imprints show

coordinates 175/049 ofthe Israeli men label refers to the grid. fragmented remains of the last abdominal tergite. Information the BMNH is The on specimens very sparse. This tergite is ornamented with paired subme- label just mentions “formation Upper Cretaceous” and “locality dian-, intermediate-, lateral-, and marginal carinae. Kerak, Moab”. Apparently these specimens originate from The submedian carinae are sharp, straight, and Senonian outcrops near the Jordanian city ofKarak in the Moab

region (Fig. 1, locality 3). Most probably the specimens were provided with a dorsal row of tubercles (Figs. 2D;

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3B). Their increases the with height gradually towards four to six tubercles, decreasing in size distally of posterior margin the tergite where they end (best in Fig. 3). The intermediate and lateral teeth The intermediate abruptly. carinae are less are Both pro- less pronounced. are inflated but not

nounced, curved, and slightly directed slightly provided with tubercles. Halfway between the oblique 2D). anterolaterally (Fig. These carinae lateral teeth and the anterior margin of the telson

to be inflated the appear medially. As is case with a wrinkle reflects the of marginal presence a pro- the submedian carinae, the intermediate carinae minent lateral crease or fold on the telson (arrows

are provided with a dorsal row of in tubercles. Lateral Figs. 2B, D; 3B, C). and carinae are in marginal only preserved GSI Between the submedian and intermediate teeth M-8113, although incompletely (Fig. 2D). The there are prominent marginal denticles. A linear lateral carinae and are straight relatively sharp. row of four equally sized submedian denticles is

These carinae also ornamented with of are a row present, arranged in symmetrical pairs (Fig. 3C).

tubercles. Thin carinae are present. These Between the submedian marginal and intermediate teeth, a carinae marginal are positioned towards linear row obliquely of four intermediate, equally sized den- the of the posterior margin tergite where they fuse ticles is present (Fig. 3C). Both submedian and with the lateral carinae. At this junction a small intermediate denticles are rounded and inflated.

posteriorly directed spine is present. Therow ofsubraedian denticles marginal is positioned more GSI M-8113 is the that shows only specimen proximally as compared to the two rows of inter-

remains of the uropods (Fig. 2C, D). The uropodal mediate denticles.

protopod structures that can be recognized include From the intermediate teeth up to the anterior

a curved lateral or a bifurcated marginal carina, edge of the telson the margin is inflated forming

median carina that is ornamented with rounded partially marginal carinae. These carinae are dor-

small dorsal tubercles, and a carina proximal pa- sally ornamented with small tubercles, on the sec- the lateral of the sixth abdominal ralleling margin tions between the anterior edge and the lateral

The 2D; seems wrinkles tergite (Figs. 4). uropodal exopod more pronounced than on the sections

of unisegmented although the presence a second between the lateral wrinkles and the intermediate

cannot be segment excluded due to the poor teeth. The sections between the lateral wrinkle and

the preservation of The is about the intermediate teeth minute uropods. exopod carry sharp, lateral

four five times as it is broad. It is to long as not tubercles on (best visible right side of Fig. 2D). to determine the overall of this possible shape The dorsal on the telson is seg- ornament very ment in full detail; of thickened distinctive only fragments (indicated in Fig. 2B). A robust rounded lateral visible. margins are The inner margin appears mediancarina is This carina the present. spans entire

inflated and carries a dorsal row ofsmall tubercles. of length the telson, starting a few millimetres from

Distally at least two lateral are The the anterior spines present. margin of the telson to end a few outer is margin more flattened and, based on the in millimetres front of the row of submedian den-

visible remains, at least the distal two third is ticles. orna- The anterior onset of the carina is rounded mented with a row of lateral and spines. slightly inflated, forming a basal notch (Figs.

Only tiny fragments of the the uropodal endopods 2B; 3B). Posteriorly carina ends gently. The

are preserved (Fig. 2D). The outer seems relative of margin sharpness the carina seems to increase

inflated, the inner carries of setae. margin a fringe posteriorly, and a median row of small tubercles The telson is about as long as broad and has the is over present its entire length, including the basal overall shape of a rectangle with convex lateral notch. The median carina is flanked by tubercles. Three margins. pairs of marginal teeth can be dis- These tubercles are organized in straight rows,

anterior tinguished: submedians, intermediates, and later- starting of the basal notch of the median als 2B; 3B). The submedian teeth the (Figs. are carina and meeting in a small arch that encloses most from the the prominent, clearly protruding pos- posterior end of the median carina. terior margin of the telson. These teeth are inflated Paired submedian carinae are situated about one their over entire and ornamented third of the length dorsally way from the median carina. These

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and of of the tubercles on the Jordanian telsons. carinae are straight slightly diverging antero- some

tubercles consist of a tubercle laterally. The carinae are only slightly elevated The seem to large

with a smaller tubercle, or simply two but are profoundly ornamented with tubercles (Fig. adjacent tubercles instead of “normal” tubercle. 3B). Posteriorly these carinae seem to form a con- smaller one

It looks like one of the endopods in tinuation of the submedian marginal teeth although uropodal

these elevated GSI, M-8113 is preserved underneath the surface there is a clear gap between both of the telson. A is visible (“un” in 2D) structures. Theanterior tip of the submediancarinae margin Fig.

A and mediad of this margin some very tiny frag- curves laterally. solitary tubercle between the

ments of a setal However, because anterior end of the submedian carinae and the an- possibly fringe. of this remarkable preservation these characters terior onset of the lateral carinae (best in Fig. 2D), should considered be- be very carefully. gives the impression of a curved connection Although the material is well preserved, fine tween these structures. lost It details were in the process of fossilization. The lateral carinae are distinctive and, as all other

could be that there are small movable or fixed apices carinae of this specimen, ornamented with a dorsal

on the submedian teeth. Especially the telsons from row of tubercles. Anteriorly the lateral carinae start remains Jordan seem to carry cuticular of very small as transverse oriented ovate notches. The carinae apical spines. However, the evidence is not con- that emanate from these notches run parallel to vincing since the rest of the teeth are not preserved the marginal carinae and end at the base of the in fine detail. submedian marginal teeth. However, these carinae The of the distal end of the presence spines on do not form continuous structures. About one third uropodal exopod is based on wet observations in from the anterior margin of the telson there is a M-8113 a smooth, circular excavation on GSI, (left fissure in the lateral carina where a low transverse side Fig. 2C). The rounded apex of the uropodal carina connects the lateral carina with the marginal exopod visible on the latex peel (right side Fig. carina. At this point there is also a low mesiad is 2D) thus a preparational artefact. protrusion of the lateral carina. This complex of

structures gives the impression of a distinctive Taphonomy. - The finds ofonly a telson with one lateral crease on the telson. Following the lateral or more attached abdominal somites are common carinae further posteriorly, there are two other low in the stomatopod fossil record (Hof & Briggs, in fissures. These fissures are positioned line with 1997). The specimens of Ursquilla yehoachi are the lateral and intermediate marginal teeth. apparently transported fragments of decayed ani-

mals. The GSI specimen carries two gastropods,

Remarks. - There is some variation in the shape one in the centre of the telson and one on the sixth from of the telsons. The telsons Jordan are abdominalsomite (Fig. 2D). These gastropods were elongated and their carinae less inflated compared apparently scavenging or grazing on the stomatopod to the Israeli telsons. However, these two differ- remains or just coincidentally fossilized on the ences could be preservational artefacts. telson. Commensal gastropods on extant stomato- carinae The number of tubercles on the various pods only attach to the abdominal or thoracic ventral also but detailed of seems to vary, a comparison side of their host (e.g. Rosewater, 1969; Reaka, the specimens is difficult because the tuberculate 1978). rows and carinae are often disturbed. On the Jor- All the specimens of U. yehoachi are no real

danian telsons the submedian teeth are ornamented imprints but external dorsal moulds. Mainly on with six dorsal tubercles, while the telsons from MNHN, R. 62691 and BMNH, 1 7316 there are Israel to four five tubercles. only seem carry or of mineralized cuticle many tiny fragments visible, The rows of tubercles flanking the median carina especially of the carinae and tubercles. Under high in the Israeli MNHN, R. 62691 count about holotype magnification (50 x) the laminated composition while 18 tubercles row the complete Jordanian per of the cuticle is clearly visible. Scanning Electron telson of BMNH, I 7316 shows about 25 tubercles. Microscope observations could reveal the fine de-

These differences are dueto the nature partly bipolar tails of these cuticular remains.

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Fig. 4. Reconstruction of the telson and the sixth abdominal tergite of Ursquilla yehoachi (Remy & Avnimelech, 1955). Of the

the visible are reconstructed; the outline is indicated uropods only fragments possible by dotted lines. Scale bar is 1 cm.

Measurements. - The length of the telson is the average length of Ursquilla yehoachi's telson, measured medially from the anterior margin to a a corse estimation of the total length would be 17

line hypothetical over the tips of the submedian or 18 cm (total length as definedby Manning, 1969). denticles. Telson width is the greatest width. The

R. 62691 3 telson of MNHN, is cm long and

3.1 cm wide. The telson of GS1, M-8113 is about Discussion

3.2 cm long and 3 cm wide; the sixth abdominal somite of this specimen is 1 cm high and about A reconstruction of the telson and the sixth

3.2 The cm broad. most complete telson on BMNH, abdominaltergite of Ursquilla yehoachi is offered

I 7316 is 2.4 cm long and estimated 2.4 cm in Fig. 4. Of the uropods only the visible remains

the wide; incomplete telson has the same dimen- are drawn. sions. The description of U. yehoachi is based on only

that Assuming these stomatopods had the body the posterior part of these stomatopods. Fortunately,

of a standard such as these of the proportions squilloid Squilla parts carry most crucial diagnostic mantis (Linnaeus, and 2.8 cm is as 1758), taken characters used in stomatopod taxonomy and a

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detailed comparison with both fossil and recent Lysiosquilla nkporoensis Forster, 1982, from the

of taxa is feasible. Upper Cretaceous Nigeria, seems to represent

The Mesozoic stomatopod fossil record en- the modernLysiosquillidae. The telson ofL. nkpo-

compasses the extinct Sculdidae Dames, 1886 and roensis is smooth and ornamented with a low

Pseudosculdidae Dames, 1886 and possible repre- mediancarina, low submedian carinae, and distinct

sentatives ofrecent stomatopod taxa. The two ex- marginal carinae. The sixth abdominal somite

tinct Mesozoic stomatopod families were lumped carries paired low, posteriorly spined submedian

by Flolthuis & Manning (1969), but should be carinae, and distinct posteriorly spined lateral

considered as clearly separate families (Hof, in carinae. Berry (1939) describedChloridella angolia

press). The major characteristic that distinguishes from the Cretaceous of Angola, a species allied to

the Mesozoic Sculdidae and Pseudosculdidae from the modem Squillidae. Unfortunately, the type

the is the of uni- material of C. imtraceable and extant stomatopods presence an angolia appears

segmented uropodal exopod. The exact structure according to Förster (1982) this species originates

of U. clear Cretaceous Eocene the uropodal exopod in yehoachi is not not from but from formations.

It the of so the systematic position of U. yehoachi should is clear that morphology of the telson

be considered on the basis of other morphological U. yehoachi is unique amongst the late Mesozoic

aspects. The general morphology of the telson fossil stomatopods. Comparison of the telson of

excludes the placement of U. yehoachi within the U. yehoachi with those of the extant stomatopods

Sculdidae. The four species of the Sculdidae all indicates this taxon can be accommodated within

have telson ornamented with of movable the The of the a a fringe superfamily Squilloidea. presence

spines and clearly lack a median carina. The telson distinct median carina on the telson and the four

of Pseudosculda laevis (Schlüter, 1874), the only intermediate denticles are diagnostic characters for

described pseudosculdid so far, possesses two the squilloids (Manning, 1995: 147). The presence

movable submedian spines and lacks the clear set of the lateral folds and the distinct tuberculate

of marginal teeth and denticles that characterize carinae are unique characters amongst the stoma-

the telson of U. yehoachi. Apart from the distinct topods and justify the erection of a new family

differences in telson morphology, it is the size of within the Squilloidea. This taxonomic placement

U. that this taxon from the that of the modern yehoachi distinguishes implies representatives super-

two extinct stomatopod families. With a length of family Squilloidea were already present by the end

around 17 or 18 cm, U. yehoachi is a real giant of the Mesozoic.

compared to the Sculdidae with their average length The value of U. yehoachi in a phylogenetic

of about 3 cm (own observations) and the slightly reconstruction of the stomatopods is unfortunately

Pseudosculdidae that between 3 limited. The and larger range to incompletely preserved uropods

cm (own observations). the lack of theentire anterior portions of the body,

Dames (1886) suggested that modem stomato- especially the second thoracopods, prevent an

pods were present at the end of the Cretaceous. exact positioning of this in the evolutionary

This remark was based solely on the record of history of the stomatopods. Within the Squilloidea,

Squilla cretacea Schluter, 1868, from the Upper however, the discovery of this fossil species might

Cretaceous ofSendenhorst in Germany. This fossil provide information useful towards polarizing

is and certain the stomatopod poorly described the accom- characters of superfamily. Some gen-

is clear. within the so-called panying drawing not very Attempts to trace era Squilloidea possess pre-

the original material have failed so far. More re- lateral lobes. These pre-lateral lobes could be

cent Mesozoic fossil stomatopod records are known remnants of the lateral folds that are so obviously

from Schram (1968) who described Paleosquilla developed in U. yehoachi. These distinct lateral

from raise brevicoxa the Cretaceous of Colombia. This folds themselves the issue of a possible two-

resemble stomatopod seems to the modern Gono- segmented origin of the modem stomatopod telson,

dactyloidea. However, the telson of this species i.e., the fusion of a telson with a seventh abdominal

was not preserved. somite. However, the Paleozoic Tyrannophontidae,

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ancestral the to modern unipeltatan stomatopods Caiman, W.T., 1904. On the classification of the Crustacea

show Malacostraca. Ann. Mag. nat. Hist., (7)13: 144-158. (Jenner et ah, 1998), no trace of such an 1886. Dames, W., Ueber einige Crustaceen aus den Kreide- amalgamation. ablagerungen des Libanon. Z. dt. geol. Ges., 38: 551-575, Lines of tubercles can be discerned on the telsons pis. XIII-XV. of modern many squilloids, and indeed the lines Forster, R., 1982. Heuschreckenkrebse (Crustacea, Stomato-

of tubercles the median carina are flanking espe- poda) aus dem Alttertiar vonHelmstedt und Handorf (Nie-

und der For dersachsen) Oberkreide von Neues Jb. cially easily recognized. example, some spe- Nigeria.

cies of the Geol. Palaont, Mh., 6; 321-335. squilloid genus Erugosquilla Manning, Glaessner, M.F., 1969. . In: R.C. Moore & C. Teichert 1995 are the of diagnosed by presence such a pattern Treatise (eds.), on invertebrate paleontology, part R, Ar- of ornament. Also extant many stomatopod spe- thropoda 4: R399-R533 (Univ. Kansas Press, Lawrence). cies outside the carry lines of tubercles Squilloidea Hof, C.H.J., 1998. Fossil stomatopods (Crustacea: Malaco-

or carinae on the dorsal face of the telson. and their J. The straca) phylogenetic impact. nat. Hist. In press.

C.HJ. & D.E.G. recently described species Faughnia formosae Hof, Briggs, 1997. Decay and mineraliza-

tion of mantis - & 1997 shrimps (Stomatopoda: Crustacea) a key Manning Chan, is an arbitrary example of to their fossil record. Palaios, 12: 420-438. a gonodactyloid with multiple dorsal carinae on Holthuis, L.B. & R.B. Manning, 1969. Stomatopoda. In: R.C. its telson. the of these Establishing homologies Teichert Moore & C. (eds.). Treatise oninvertebrate pale- structures with the noted for U. pattern yehoachi ontology, part R, Arthropoda 4: R535-R552 (Univ. Kan-

would detailed sas require a very comparative study. Press, Lawrence).

If U. indeedbe in the Jenner, R.A., C.HJ, Hof & F.R. Schram, 1998. Palaeo- and yehoachi can placed super- archaeostomatopods (Hoplocarida, Crustacea) from the Bear family Squilloidea, and if P. brevicoxa Schram, Gulch Limestone, Mississippian (Namurian), of central 1968 and L. nkporoensis Forster, 1982 represent Montana, Contrib. ZooL, Amsterdam, 67: 155-185. respectively the superfamilies Gonodactyloidea and Latreille, P.A., 1803. Histoire naturelle, generale et particu- this would that the three Lysiosquilloidea, mean liere, des Crustaces et des Insectes, 3: 1-468 (F. Dufart, largest superfamilies of the modern stomatopods Paris).

1806. Genera Crustaceorum et Latreille, P.A., Insectorum se- were definitely present at the end ofthe Mesozoic. cundum ordinem naturalem in familias disposita, iconibus U. yehoachi perhaps does not yet provide the ulti- exemplisque plurimis explicata, 1: 1-302 (Amand Koe- mate information to reconstruct the origin of the nig, Parisiis & Argentorati). modern stomatopods, but it is certainly another Latreille, P.A., 1817. Les Crustaces, les Arachnides et les

in the of the Insectes. In: G. Cuvier Le interesting piece evolutionary jig-saw (ed.), regne animal distribue

d’apres son stomatopods. organisation, pour servir de base a 1’histoire

et naturelle des animaux d’introduction a 1’anatomie

comparee. Ed. 1, 3: i-xxix, 1-653 (Deterville, Paris).

naturelles du Latreille, P.A., 1825. Families regne animal, Acknowledgements et ordre exposees succinctement dans un analytique, avec

1’indication de leurs 1-570 genres: (J.B. Bailliere, Paris).

For me with the I wish to L. C., 1758, tria providing specimens thank Agnes Linnaeus, Systema naturae per regna naturae,

de MNHN, David N. Lewis secundum charac- Rage (Institut Paleontologie, Paris), classes, ordines, genera, species, cum

(Collections Manager, Fossil Invertebrates and Plants, De- teribus, differentiis, synonymis, locis, Ed. 10, 1: i-iii, 1-

of partment Palaeontology, BMNH, London), and Zeev Lewy 824 (Laurentii Salvii, Holmiae).

(GSI, Jerusalem). ZeevLewy also helped me with the description Manning, R.B., 1969. Stomatopod Crustacea of the Western

ofthe localities and the stratigraphical data. I thank John Jagt Atlantic. Stud. trop. Oceanogr., 8: i-viii, 1-380 (Miami

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imen to Amsterdam. Frederick R. Schram and Lipke B, Holt- Manning, R.B., 1995. Stomatopod Crustacea of Vietnam: The

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