NovitactesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3205, 9 pp., 2 figures August 29, 1997

A Pulicid in Miocene Amber from the Dominican Republic (Insecta: Siphonaptera: ) *

ROBERT E. LEWIS1 AND DAVID GRIMALDI2

ABSTRACT A new of flea, larimerius, is de- its is found throughout the Western Hemi- scribed on the basis of a complete female speci- sphere and contains one species known to be a men in Miocene amber from the Dominican Re- cosmopolitan pest. The six described, extant spe- public. Although at least eight other, exceptionally cies of Pulex feed on a great variety of mamma- rare are known from Dominican amber, the lian hosts, and even a species of bird. The pres- ones that have been studied all belong to the Rho- ence of five strands of hair in the amber with the palopsyllidae. The specimen described here is flea virtually confirms the mammalian host of the unique as the only known fossil in the Pulicidae; fossil.

INTRODUCTION Amber is renowned for the exquisite pres- ment in tree resin: midges that swarmed too ervation of small organisms that became en- close to the tree trunk, ants that scurried trapped in it millions of years ago when it along the tree trunk, stingless bees that har- was sticky resin. Most life forms in amber vested the resin for their nests. As one might had habits (based on inference from modern expect, ectoparasitic are extreme- relatives) for which we would expect entrap- ly rare in amber, as they are in any fossil

* Journal Paper No. J-16752 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa; Project 3100. I Professor Emeritus, Department of Entomology, Iowa State University, Ames, IA 50011-3222. Present Address: 2906 Stone Brooke Circle, Ames, IA 50010-4174. 2Chairman and Curator, Department of Entomology, American Museum of Natural History.

Copyright X American Museum of Natural History 1997 ISSN 0003-0082 / Price $1.60 2 AMERICAN MUSEUM NOVITATES NO. 3205 deposit. We report one such exceptional fos- in the Staatlichmuseum fur Naturkunde, sil here, in lower Miocene amber from the Stuttgart (photo in Grimaldi, 1996: 115) and Dominican Republic. the other three in private collections. These Fossil fleas, or presumed fleas, have been appear to be rhopalopsyllids as well. Poinar previously reported from Baltic amber as (1992) reported, based on unpublished work well as several sedimentary deposits. Pa- by Traub, that even though most extant rho- laeopsylla klebsiana Dampf, 1911, and Pa- palopsyllids feed on rodents and marsupials, laeopsylla dissimilis Peus, 1968, were de- morphological features of the three speci- scribed from Baltic amber. According to mens studied by Traub indicate that they Larsson (1978) and Poinar (1992, 1995b), were bird parasites. The only living members these two species belong to the family Hys- of this family that are avian ectoparasites be- trichopsyllidae, but Palaeopsylla, a primitive long to the genus Parapsyllus, most of which genus with 49 living species, is now assigned are found on burrow-nesting sea birds in the to the hystrichopsylloid family Ctenophthal- Southern Hemisphere. A few specimens of midae. The species in this genus are mainly Listronius robertsianus (Jordan, 1938) have parasites of insectivores and are restricted to been taken from penguin nests in the Falk- the Palearctic Region. Fleas in Baltic and land Islands. Poinar (1995b) later changed Dominican amber are the only known Terti- his view, arguing that the Dominican amber ary fossils, although there are five Creta- rhopalopsyllids had capromyid rodent hosts. ceous taxa that are presumptive fleas or per- Birds did exist in the Dominican amber haps closely related to the order Siphonap- forests, and various rare feather inclusions tera (Lewis and Grimaldi, in prep.). have been reported by Poinar (1992) and Dominican amber is known for the great Grimaldi (1996). In fact, one feather in the variety of organisms preserved in it, perhaps Smithsonian collection has been identified, even more diverse than the Oligocene/Eo- on the basis of microscopic features of the cene Baltic amber that has been studied for barbules, to be from a piculet woodpecker (Layboume et al., 1994). centuries (Larsson, 1978). The age of Do- An incredible aspect of the piece of amber minican amber has been unnecessarily con- we are reporting on here is the presence of fused. An Oligocene and Eocene age (30-40 five strands of fine hair, obviously of mam- Ma) has been given by Poinar (1992, 1995a, malian origin. Although this is not definitive 1995b), but stratigraphic information indi- proof that the hair came from the host of the cates that this amber is actually Miocene flea, the conclusion is unavoidable, based on (Grimaldi, 1995; Iturralde-Vinent and the taxonomic position of the fossil and the MacPhee, 1996). Fossils belonging to 24 or- probability of its preservation. Fleas are ex- ders of are known from Dominican tremely rare in Dominican amber, with only amber, including such small and rare orders eight specimens known from approximately as Zoraptera (Poinar, 1992; Grimaldi, un- 300,000 pieces of amber (1: 30,000 or 1:3 publ.). No mallophagan or anopluran lice X 104). Hair is also very rare in amber. One have been found thus far, nor have bird and of us (Grimaldi) has found only three pieces bat flies (Hippoboscidae, Nycteribiidae, and containing hair in approximately 100,000 Streblidae), but extremely rare ticks occur in pieces of amber screened over the last ten this amber (Poinar, 1992; Grimaldi, 1996) years (1: 30,000 or 1: 3 X 104). The likeli- and even rarer are the fleas. hood of a flea and hair strands being captured Poinar (1992) mentioned three fleas stud- simultaneously, by chance alone, is incon- ied by R. Traub, all of which are reported to ceivably remote: approximately 1 in 108. belong to the Rhopalopsyllidae; two in the Surely this fossil flea fed on a mammal, Traub collection, the other in the United which would make it the only known fossil States National Museum, Smithsonian Insti- flea preserved with remnants of its host. tution. Since then, Poinar reported (1995b) another rhopalopsyllid, identified by R. E. SYSTEMATICS Lewis as Rhopalopsyllus sp. That specimen Genus Pulex is in Poinar's personal collection. There are Pulex. Linnaeus, 1758. Syst. Nat., 10th ed., I: 614 at least four more unstudied specimens: one (in Europa, America). 1997 LEWIS AND GRIMALDI: FLEA IN AMBER 3

TYPE SPECIES: P. irritans Linnaeus, 1758, not until 1958 that the name was finally res- by subsequent designation by Baker, 1904. urrected by Smit. From 1908 to 1958 all GENERIC DIAGNOSIS: Anterior margin of North American collections were assigned to head smoothly rounded or slightly angled, P. irritans. Though the two species are prac- frontal tubercle absent or inconspicuous. Eye tically impossible to separate in the females present, reduced in P. sinoculus, never sin- and differences in the males are virtually uate or with an internal sinus. Antennal cla- cryptic, they do show some differences in vus asymmetrical, its dorsal margin folea- host preferences that allow us to at least cious. Labial palpi four-segmented. Some speculate about the true nature of the records species with vestigial genal comb of one or reported during the 50 years that P. simulans two spines per side. Pronotal comb absent. was not acknowledged to be a valid species. Mesothoracic pleural rod absent. Metanotum First, P. simulans is restricted to the West- and first abdominal tergite of about equal ern Hemisphere where it is known from Flor- length dorsally, or the former slightly longer ida, west and north to southern Alberta and than the latter. Spiracle of metepimeron larg- British Columbia in North America, south to er than abdominal spiracles. Tarsal segment the northern half of Brazil and northern Chile V of all legs with four pairs of lateral plantar in South America. Haas and Wilson (1967) bristles, the space between pairs three and reported this species from Hawaii. It parasit- four wider than that between pairs one and izes a broad range of hosts (Didelphis [Di- two and two and three. Inner surface of hind- delphimorphia], Canis, Mephitis, and Uro- coxae with row or patch of short, spiniform cyon [Carnivora] Spermophilus [Rodentia], bristles. Sensilium with 14 pits per side. Bul- and Odocoileus [Artiodactyla]) but it seems ga of female spermatheca globular. Not all of to be primarily a parasite of prairie dogs (Cy- these characters are visible in the fossil spec- nomys [Rodentia]) and other burrow-dwell- imen. ing hosts. It occurs at the lower elevations in COMMENTS: The flea specimen we describe Central and South America and is replaced here is unusual in that it not only belongs to by P. irritans at higher altitudes, as pointed the Pulicidae, it is also a member of an extant out by Hopla (1980), but sufficient material genus: Pulex. This genus is indigenous to the to analyze the distribution is not available at Western Hemisphere, with one species, P. ir- this time. Both Hopla (1980) and Smit ([for- ritans, found throughout the world. It is merly curator of Siphonaptera at the British sometimes divided into two subgenera, each Museum of Natural History] personal com- with three species. The species of (Juxtapu- mun. to Lewis) are of the opinion that this is lex) are P. alvarezi Barrera, 1955, from the a complex genus and that there are a number Central American tapir (Perissodactyla); P. of undescribed South American species al- echidnophagoides (Wagner, 1933) from lied to P. simulans that must be studied be- southern Costa Rica from armadillos (Xe- fore it can be accurately delineated. narthra), and P. porcinus Jordan and Roths- Pulex irritans is also certainly native to the child, 1923, from southern Texas and Mexico Western Hemisphere and there it is sympatric from peccaries (Artiodactyla). with P. simulans except in much of north- The nominate subgenus, to which this fos- eastern North America and southern South sil species is assigned because it lacks an an- America where P. simulans has not been gular frons or any sign of a frontal tubercle taken. Pulex irritans has been transported to (characters found in members of the other other parts of the world via human agencies, subgenus), contains P. sinoculus Traub, but P. simulans evidently has not. In some 1950, from Guatemala from pocket gophers parts of the world P. irritans is most com- (Rodentia) and the P. irritans/simulans com- mon on large, hole-dwelling hosts such as plex. The taxonomic history of P. irritans foxes, porcupines, and badgers, and some- and P. simulans is somewhat confused. Until times it becomes a pest of domestic . Baker erected P. simulans in 1895, all North While definitely a parasite of mammals, a American records were assigned to P. irri- population of this species parasitizes the bur- tans. Jordan and Rothschild (1908) synony- rowing owl (Speotyto cunicularia) from mized P. simulans with P. irritans and it was southem British Columbia to northem Utah, 4 AMERICAN MUSEUM NOVITATES NO. 3205

r.w 400*1~ ~ ~ fAZ

Fig. 1. Photograph of AMNH DR-14-1140, showing most inclusions, particularly the strands of hair (arrows) and the flea (arrow with F). and western Montana and Wyoming to east- present and conspicuous, but somewhat re- ern Washington and Oregon. Hopla (1980) duced in diameter, resembling that of P. sin- found the species to be remarkably homo- oculus. Labial palpi stiff and sclerotized, geneous morphologically in spite of its broad their basal segmentation obscured. Mesotho- distribution. However, because of its distri- rax without internal pleural rods. Metanotum bution, it has been reported from a multitude slightly longer dorsally than abdominal ter- of hosts, most of them probably accidental gum I. Tarsal segment V of all legs with four associations. pairs of lateral plantar setae, the distal pair Hopla (1980) discussed the genus in detail, somewhat more delicate than the proximal especially the host associations and zooge- three pairs. Tarsal claws of all legs long, ography of the Pulex irritans/simulans com- slender, and bearing a slender basal tooth. All plex. abdominal tergites and most sternites with a single row of long setae. No visible tergal Pulex (Pulex) larimerius, new species spinelets. One long antesensilial seta per Figures 1, 2 side. Sensilium small, convex, and with a HOLOTYPE: Female, AMNH DR-14-1140. well-developed anterior collar. ETYMOLOGY: Named in honor of Roy and DESCRIPTION: Body length ~-2.19 mm. Maria Larimer, who donated the specimen to Head. Head capsule smoothly rounded, its the AMNH because of its scientific value. frontal and occipital margins separated by DIAGNOSIS: Anterior margin of head narrow but well-defined falx. Frontal tuber- smoothly rounded and lacking a frontal tu- cle absent. Eye well developed but somewhat bercle. Interantennal suture (falx) present. reduced in diameter compared to other mem- Antennal clavus asymmetrical (though miss- bers of the subgenus except P. sinoculus. ing from left side and distorted on right). Eye Three setae in ocular row but no evidence of 1997 LEWIS AND GRIMALDI: FLEA IN AMBER 5

Fig. 2. Top: Photomicrograph of flea in AMNH DR-14-1140. Bottom: Illustration of flea. 6 AMERICAN MUSEUM NOVITATES NO. 3205 a genal spine. Antennal fossa deep, one side surface. Longest seta in apical notch extend- containing a shriveled antenna with an asym- ing about 3/4 distance to apex of hindtarsal metrical clavus. One long seta per side about segment I. Hindtarsal segment IV hardly as midway along the dorsal margin of the an- long as wide, segment III slightly longer than tennal fossa. Both frontal and occipital areas wide, segments II and V about equal in of head capsule punctate. Occipital setal row length and segment I the longest. The longest of three long bristles per side, not extending apical seta of segment I extends beyond the over dorsal margin of head. Maxillary lobe apex of segment II, that of segment III ex- triangular, its palpus four-segmented. Labial tends beyond the apex of segment IV, and palpi stiff, sclerotized, their basal segmenta- segment V bears the chaetotaxy as described tion obscured, but with a least four visible for the other legs. Unmodified Abdominal segments, the terminal segment about twice Segments. Abdominal tergum I with four se- the length of the other visible segments. tae per side in a single row. Terga II-VI with Thorax. Pronotum with five or six setae per five or six setae per side, also in a single row. side in a single continuous row over dorsum. Tergum VII with one row of about four long Prosternosome bare. Mesonotum also with a setae and one antesensilial seta per side on continuous row of five or six long setae per its caudal margin. Tergum VIII with about side. Mesepisternum with two, and mesepi- 12 setae per side on its outer surface and an- meron with four long setae per side. Meta- other 8 to 10 setae per side along its caudal notum with six long setae per side preceded margin. Sterna III-VII with a single row of by two irregular rows of shorter bristles but four to six setae per side. Caudal margin of lacking marginal spinelets. Metepisternum sternum VII slightly concave and sternum bare, but metepimeron with ten long bristles VIII triangular. Genital Segments. Sensilium arranged in two irregular rows of five each. small, slightly convex dorsally, with a well- Legs. Outer surface of forecoxa with a num- developed anterior collar, but the number of ber of oblique rows of setae, most of which pits cannot be determined (14 is normal for are missing. Foretrochanter and forefemur the family). The anal stylets, spermatheca, distinct but bare. Foretibia with about four and genital ducts are not visible. notches in dorsal margin bearing pairs of long setae, the longest in apical notch ex- REMARKS tending about half way to the apex of fore- tarsal segment II. All tarsi five-segmented. Fleas are usually described from material Foretarsal segments I and IV shortest, about mounted on microscope slides and are thus as long as wide, segment III slightly longer, only examined in one aspect. However, am- segment II still longer, and segment V the ber fossils can be manipulated in ways that longest, with four visible pairs of lateral reveal structures in more than one aspect. plantar setae, the apical pair much smaller Therefore, it should be understood that the than the other three pairs and spaced as is accompanying drawing is a composite of typical for the genus. Foretarsus terminated both sides of the specimen rather than an il- by pair of long, thin, tarsal claws (ungues) lustration of one side. each with a long, thin, basal tooth. Dorsal The amber in which the specimen is pre- margin of midtibia with five notches bearing served is a flat, oval piece, 21 mm long X paired, long setae. Midtarsal segments sub- 12 mm wide and 4 mm thick, containing the equal in length, segments I and IV slightly following inclusions: the flea, tiny pyritic longer than wide, segment III about twice as crystals ('20 ,um diameter) scattered long as wide, segment II about three times throughout two fracture planes that parallel as long as wide, and segment V slightly lon- the surface (the flea is immediately under one ger, with chaetotaxy and ungues as described of these planes); cleared/skeletonized phorid for foretarsus. Hindcoxae with three or four fly; decayed, exoskeletal remains of uniden- mildly spiniform bristles apically on inner tifiable insects; two very tiny (0.35 mm) surface. Dorsal margin of hindtibia with five wingless hexapods, probably neanurid or on- notches bearing paired long setae, as well as ychiurid collembolans (a furcula is absent); a submarginal row of six fine setae on outer a nymphal oecanthine (tree) cricket; several 1997 LEWIS AND GRIMALDI: FLEA IN AMBER 7 fecal pellets and particulate debris; and the family Atopomelidae suggests that the five strands of fine hair (the longest and most hairs belonged to a rodent. The bulk of the complete is 9.65 mm long and 50 p,m thick). New World, Northern Hemisphere species of these mites are parasites of capromyid ro- COMMENTS dents, although one is known from a species of Solenodon, and others from New World Because this flea belongs to an extant ge- monkeys that are now extinct in the Carib- nus, and the piece of amber has hair in it, bean basin (B. M. O'Connor, personal com- questions of the authenticity of the amber mun.). The amber described in the other re- may arise. The piece is unquestionably au- port (Poinar and Columbus, 1992) contains thentic Dominican amber, based on a variety 40 strands of hair and an adhesive grass of features and circumstances. The other in- spikelet to which one of the strands is at- clusions in the amber, their state of preser- tached. The cuticular pattern of the hair sug- vation, the fact that a hot needle touched to gested to these authors that it came from a the surface produces a resinous smell, not carnivore. Poinar (1992) mentioned that al- like burning plastic, and that the piece fluo- though some mammals (certain bats and resces under long-wave UV light, all attest sloths) have distinctive hair structure, iden- to its authenticity. Also, the piece was dis- tification of hair in amber is problematic. In- covered among a group of approximately deed, variation among hairs from the same 1000 small pieces of Dominican amber and individual of many mammals is known to en- was not being sold under any pretense. compass the variation seen among various mammalian orders (Brunner and Coman, DISCUSSION 1974; Debrot et al., 1982). Recently, ex- MAMMALS CONTEMPORANEOUS WITH tremely well-preserved hair in late Paleocene DOMINICAN AMBER coprolites from Mongolia was reported by Meng and Wyss (1996). The hairs were pre- The host of this flea was certainly mam- served with the skeletal remains of the mul- malian. There are several other specimens of tituberculate, Lambdopsalis, to which they Dominican amber containing hematophagous presumably belonged. This hair is very sim- vertebrate parasites preserved with remains ilar to that of modern mammals, indicating of their host. Piece number DR-12-857 in the that evolutionary changes in the fine struc- AMNH contains a tick belonging to the ixo- ture of mammal hair is extremely conserva- did genus Amblyomma with two complete tive. The multituberculates are the most strands of short, fine, stiff hair. There is a primitive clade of extinct mammals and orig- remarkable specimen in the Stuttgart collec- inated in the Triassic. tion (DO-5514-M) containing a small swarm The best evidence for a mammal contem- of phlebotomine sandflies with approximate- poraneous with Dominican amber was re- ly 10 strands of hair and debris, probably ported by MacPhee and Grimaldi (1996). from a mammal nest. Most of the female Skeletal remains of an insectivore were sandflies are replete, probably from a blood found in a piece of amber recovered directly meal. A photograph of a portion of the piece from the La Toca amber mines near Santiago, is shown in Grimaldi (1996: 114). Micro- Dominican Republic, by Grimaldi. The spec- scopic features of the hairs in these two piec- imen, AMNH DR-14-627, consists of six es have yet to be studied. An argasid tick was thoracic vertebrae with portions of four ribs reported in a piece of Dominican amber with still articulated. The vertebrae are very sim- a probable mammalian scat (Poinar, 1995a). ilar to those of certain insectivores such as Hair has been previously reported from the Caribbean endemic solenodontids, Solen- Dominican and even Mexican amber (Poinar, odon and Nesophontes. Unfortunately, no 1988, 1992; Poinar and Columbus, 1992). hair is preserved in the piece of amber. One piece from Costa Collection in Puerta Despite claims that "no findings of fossil Plata, Dominican Republic (plate 3 in Poinar, mammals older than 10,000 BP [occur] in 1992) contains a clump of numerous fine Hispaniola" (Poinar, 1995b), fossil evidence hairs. The presence of a mite belonging to does exist for Miocene land mammals in 8 AMERICAN MUSEUM NOVITATES NO. 3205

Puerto Rico and Cuba, and thus they must species are remarkably similar in their mor- have occurred in Hispaniola as well. The phology, in spite of their diverse host pref- mammals are a megalonychid sloth from the erences. This is particularly the case in the Miocene of Cuba (and another possible one females, which show only cryptic differences from the Oligocene of Puerto Rico), and a between the species. Although internal struc- capromyid rodent and platyrrhine primate, tures such as the genital ducts and sperma- also from the Miocene of Cuba (MacPhee theca cannot be seen in this fossil, its exter- and Iturralde-Vinent, 1994, 1995). nal morphology is remarkable for its simi- larity to females of living species, indicating SYSTEMATIC AND EVOLUTIONARY that little morphological evolution has taken SIGNIFICANCE OF A LOWER MIOCENE PULICID place in the genus since the Miocene. (Whether this is viewed as a primitive or As indicated earlier, our understanding of de- the composition of the genus Pulex is far rived condition is a matter for conjecture.) from complete and large collections of ma- As noted, all of the fossil fleas previously terial from Central and South America are known to date from Dominican amber have required before the alpha of this belonged to the predominantly Neotropical genus can be thoroughly known. Material to family Rhopalopsyllidae, most members of support such a study is not presently avail- which are parasites of small rodents. The or- able. However, the Pulicidae is thought by igin and New World distribution of species most pulicologists to be an advanced family of Pulex certainly would favor the presence compared to most others in the order and of pulicid fleas in Dominican amber. Their quite distinct, in spite of the diversity reflect- host preferences for larger animals probably ed in its current subfamily classification. Pu- prevented them from coming into contact lex is the type genus for both the Pulicinae with the plant resins from which amber ul- and the tribe , and the six recognized timately developed.

REFERENCES Baker, C. F in Hawaii (Siphonaptera: Pulicidae). J. 1895. Preliminary studies in Siphonaptera. II. Med. Entomol. 8: 25-30. Family Pulicidae Tschb. Can. Entomol. Hopla, C. E. 27: 63-66. 1980. A study of the host associations and zo- 1904. A revision of American Siphonaptera, ogeography of Pulex. In R. Traub and or fleas, together with a complete list H. Starke (eds.), Fleas. Proc. Int. Conf. and bibliography of the group. Proc. Fleas. Peterborough, UK: Ashton U.S. Natl. Mus. 27: 365-469. Wold, pp. 21-25, June 1997. Rotter- Brunner, H., and B. Coman dam: A.A. Balkema, 1980. 1974. The identification of mammalian hair. Iturralde-Vinent, M. A., and R. D. E. MacPhee Melbourne: Inkata Press. 176 pp. 1996. Age and palaeogeographical origin of Debrot, S., G. Faviz, C. Mermod, and J.-M. We- Dominican amber. Science 273: 1850- ber 1852. 1982. Atlas des poils de mammifres d'Eu- and N. C. Rothschild rope. Peseux, Suisse: Imprimerie de Jordan, K., l'Quest S.A. 208 pp. 1908. Revision of the non-combed eyed Si- Grimaldi, D. phonaptera. Parasitology 1: 1-100. 1995. The age of Dominican amber. In K. B. Larsson, S. G. Anderson and J. C. Crelling (eds.), 1978. Baltic amber: a paleobiological study. Amber, resinite and fossil resins, pp. Entomonograph 1. Klampenborg, Den- 203-217. Washington, D. C.: American mark. Chemical Society Symposium 617. Laybourne, R. C., D. W. Deedrick, and E M. Hue- 1996. Amber: window to the past. New York: ber Abrams/Am. Mus. Nat. Hist. 1994. Feather in amber is earliest New World Haas, G. E., and N. Wilson fossil of Picidae. Wilson Bull. 106: 1967. Pulex simulans and P. irritans on dogs 18-25. 1997 LEWIS AND GRIMALDI: FLEA IN AMBER

Lewis, R. E., and D. Grimaldi land mammals In prep. The tertiary [sic!] in the An- status of fossil Siphonaptera. tilles. Experientia 44: 88-89. MacPhee, R. D. E., and M. A. Iturralde-Vinent 1992. 1994. First Life in amber. Palo Alto: Stanford Tertiary land mammal from Great- Univ. Press. er Antilles: an early Miocene sloth (Xe- narthra, Megalonychidae) from 1995a. First fossil soft ticks, Ornithodoros antiq- Cuba. uus n. Am. Mus. Novitates 3094: 13 pp. sp. (Acari: Argasidae) in Domin- 1995. Origin of the Greater Antillean land ican amber, with evidence of their mam- mammal fauna, 1: New Tertiary fossils malian host. Experientia 51: 384-387. from Cuba and Puerto Rico. Am. Mus. 1995b. Fleas (Insecta: Siphonaptera) in Do- Novitates 3041: 31 pages. minican amber. Med. Sci. Res. 23: 789. MacPhee, R. D. E., and D. Grimaldi Poinar, G. O., Jr., and J. T. Columbus 1996. Mammal bones in Dominican amber. 1992. Adhesive grass spikelet with mamma- Nature 380: 489-490. lian hair in Dominican amber: first fos- Meng, J., and A. R. Wyss sil evidence of epizoochory. Experien- 1996. The first report of multituberculate hair tia 48: 906-908. and a reappraisal of auditory anatomy Smit, F G. A. M. of Lambdopsalis. J. Vertebr. Paleontol. 1958. A preliminary note on the occurrence (in press). of Pulex irritans L. and G. Pulex simulans Poinar, O., Jr. Baker in North America. J. Parasitol. 1988. Hair in Dominican amber: evidence for 44: 523-526. w

Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last five years are available at World Wide Web site http://nimidi.amnh.org. Or address mail orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024. TEL: (212) 769-5545. FAX: (212) 769-5009. E-MAIL: [email protected]

0 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).