AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3140, 7 pp., 17 figures June 29, 1995

Caddis Flies (Trichoptera) from Turonian (Upper ) Amber of New Jersey1

LAZARE BOTOSANEANU2

ABSTRACT Three well-preserved caddis flies were discov- small representative ofAgraylea may be the oldest ered in two pieces ofTuronian-age amber (Upper known hydroptilid. The third specimen is the male Cretaceous, 90-94 million years old [Ma] from of a species of the Recent, Miocene, Oligocene, central New Jersey. Two ofthem are the male and and Upper Cretaceous philopotamid genus Wor- female of a new species of the Recent and Oligo- maldia, being the oldest known species certainly cene hydroptilid genusAgraylea (sensu lato); a new belonging to this genus. subgenus is described for this species; this very

INTRODUCTION The American Museum ofNatural History to large pieces 10-15 cm long and 8-12 cm is excavating amber from an unexpectedly wide. The substance is apparently the fossil- rich deposit from the Raritan-Magothy For- ized resin of araucarian trees, based on the mation (Upper Cretaceous: Turonian, ca. 90- structure of large pieces of carbonized wood 95 Ma) of central New Jersey. Grimaldi et found with the amber as well as on the chem- al. (1989) summarized the paleontology of istry ofthe amber. The site ofthe excavations New Jersey fossil resins. The amber can vary is undoubtedly a lagoonal or deltaic deposit, from transparent yellow to red, turbid, and with carbonized wood, conifer needles, stems cloudy, from pieces only 2-3 mm in diameter ofEquisetales, and the amber associated with

I Paleontology of New Jersey Amber, Part III. 2 Honorary Research Fellow, Zoological Museum (Entomology), Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands.

Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $1.40 2 AMERICAN MUSEUM NOVITATES NO. 3140 them, representing stranded, redeposited ma- Specimen generally well preserved. Head with terial. ocelli and two distinct pairs of large, setose Among the hundreds ofamber pieces con- warts (the presence of a third pair is not ex- taining inclusions, two were found cluded). Antenna with 22 segments, typically with Trichoptera, described below. hydroptilid, with fine, short setae. Maxillary palpi cannot be well observed, whereas a la- bial palpus is represented in figure 1. No in- ACyKNOWLEDGMENTS teresting details of the thorax could be ob- Thanks are expressed to David Grimaldi served. Leg spurs clearly distinct: 1, 3, 4 (fore- (AMNH) for providing this interesting ma- leg spur well developed; internal spurs much terial plus photographs, for information on longer than external spurs). Left forewing en- the amber and the site, as well as for editing tirely distinct, but with tip slightly twisted; the manuscript. Thanks are also due to Ge- right forewing not as well preserved, never- rard R. Case, James J. Leggett, and Paul D. theless with many details distinct; forewings Borodin, for their collecting efforts. broad, apex not pointed but almost parabol- ic, setose, with moderately long fringes and thickened setae along SC vein. SC anteapi- SYSTEMATICS cally connected by a transverse vein to RI; FAMILY HYDROPTILIDAE f1, f2, and f3 present; fi distinctly shorter (and wider) than f2; also f3 shorter than f2, but Genus Agraylea Curtis, 1834 longer than fi and narrow; transverse vein DIAGNOSIS: (s.l., includingAllotrichia McL., connecting RS with M distal to bifurcation 1880): Mostly relatively large Hydroptilidae; of M; positions of A2, A3, and A4 rather cu- wings relatively broad, apex parabolic; ve- rious; A4joining A2, and A3 joining A4 slight- nation fairly complete (f 1, 2, 3 in forewing; ly more proximally (fig. 2). Hindwings less 1,2, 3, 5 or 2, 3, 5 in hindwing); ocelli present; well suited for observation, but essential de- tibial spurs 0, 3, 4 (only in some extinct spe- tails are distinct: not strongly pointed; SC cies 1, 3, 4). joins RI at level with bifurcation of RS; fi short, distinct in right wing, absent in left; f2, Nanoagraylea, new subgenus f3, f5 present. Last abdominal segments are very well preserved, allowing examination of DIAGNOSIS: Very small size: in recent their ventral side; sternite VIII without any Agraylea s.l., the wing expanse is mostly 8- distinct mediolongitudinal formation, api- 11 mm, 6-8.5 mm in the Oligocene species, cally with crown of 10 long setae. and only 4-5 mm in the new Cretaceous spe- MALE: Far less well preserved than the fe- cies; very short fl in forewings; fl in hind- male. Forewing length about 2.2 mm (wing wings present or absent; no median process expanse thus about 4.9 mm). General color on male sternite VII; no particular formation ofbody and wings dark brown. Head offering on female sternite VIII. no clear and relevant details. Antenna with TYPE SPECIES: Agraylea cretaria, n. sp. 26 segments. Spurs exactly as in female. Wings badly preserved, but nothing in the observ- Agraylea cretaria, new species able details casts doubt about this specimen (Figures 1-8, 14-16) belonging to the same species as the female TYPES: Female holotype and male allotype (f1, f2, and f3 can be well observed in the right in the same piece of yellow amber from forewing, and less well in the left one). Apex Sayreville, Middlesex Co., New Jersey, G. R. of abdomen strongly twisted and partly Case, coll. AMNH NJ-86. Despite the fact opaque; nevertheless, a fairly good dorsoap- that the male is somewhat larger than the ical view of the genitalia was obtained. A female, I have no doubts about the fact that median process on sternite VII is definitely they belong to the same species. absent. Segment X very deeply (maybe com- DEsCRIPTIoN OF FEMALE: Forewing length pletely) split medially, the internal borders of 1.8 mm, wingspread thus about 4 mm. Gen- its two halves deeply notched in a very char- eral color of body and wings dark brown. acteristic manner; gonopods elongate, rather 1995 BOTOSANEANU: TRICHOPTERA FROM TURONIAN AMBER 3

5 Figs. 1-5. Agraylea (Nanoagraylea, n. sg.) cretaria, n. sp., female holotype. 1: Head, lateral. 2, 3: Left forewing and distal parts of right forewing. 4, 5. right and left hindwings. narrow, apparently of simple shape and api- nus surviving, one species from Baltic Oli- cally thickened; long tufts ofstrong setae arise gocene amber); Allotrichia McLachlan, 1880 from below, probably inserted on sternite IX. (two species from Baltic Oligocene amber; ETYMoLoGY: Subgenus: Nanus (L.) = dwarf; Allotrichia is considered here only as a sur- species: cretarius (L.) = belonging to chalk viving subgenus of Agraylea); Electrotrichia (an early name for the Cretaceous). Ulmer, 1912 (genus extinct, one species from DIscussIoN: At present there are 15 species Baltic Oligocene amber); Leucotrichia Mose- belonging to the family Hydroptilidae Ste- ly, 1934 (genus surviving, one species from phens, 1836, that have been described from Early Miocene Dominican amber); Ochrotri- fossil material. These species were described chia (sg. Ochrotrichia) Mosely, 1934 (genus in the following genera (here chronologically surviving; 6 species-one ofthem not named, arranged): Burminoptila Botosaneanu, 1981 one still surviving-from Early Miocene Do- (genus extinct, one species in minican amber, but age ofone ofthem given of unknown age, but probably lower Terti- as "Oligocene"); Alisotrichia Flint, 1964 (ge- ary); Hydroptila Dalman, 1819 (genus sur- nus surviving, one species from Early Mio- viving; one species from Eocene ofColorado cene Dominican amber). not surely belonging to this genus or even to Agraylea (Nanoagraylea) cretaria, n. sp., is the family); Palaeagapetus Ulmer, 1912 (ge- probably the oldest known representative of nus surviving, with one species from Baltic the Hydroptilidae; there is a hydroptilid from Oligocene amber); Agraylea Curtis, 1834 (ge- Burmese amber (see above), unfortunately of 4 AMERICAN MUSEUM NOVITATES NO. 3140

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Figs. 6-8. Agraylea (Nanoagraylea, n. sg.) cretaria, n. sp. 6. Terminalia of female holotype, ventral. 7. Antenna of male allotype. 8. Genitalia of male allotype, dorsoapical view. uncertain age (Tertiary or Upper Cretaceous) idence, incomplete as it may be, is additional (Botosaneanu, 1981). evidence that Allotrichia cannot be consid- The new species certainly belongs, by all ered a distinct genus from Agraylea. genitalic and nongenitalic characters, to the It is interesting to note that the new species Palearctic and Nearctic genus Agraylea (sen- shares with one of the Oligocene species, su lato, i.e., including Allotrichia: see Mar- Agraylea (Allotrichia) succinica, a well-de- shall, 1979: 193, 197; Botosaneanu, 1992: veloped spur on the forelegs, which is lost in 50). Three species ofAgraylea and Allotrichia all other known species of Agraylea s.l., ex- were very well described or redescribed from tinct or recent. Oligocene Baltic amber by Ulmer (1912): Agraylea spathifera Ulmer, 1912; Allotrichia FAMILY ampullata Ulmer, 1912; and Allotrichia suc- Genus Wormaldia MacLachlan, 1865 cinica Hagen, 1856. The new species de- scribed here from New Jersey amber (Turo- DIAGNosIs: Medium-sized Philopotami- nian, 90-94 Ma) is about 50 m.y. older, but dae; 2nd segment of maxillary palpi short; shares with the Oligocene Allotrichia species tibial spurs 2, 4, 4; tibiae and tarsi of legs of an fi in the forewings shorter than f2; in one female not dilated; inferior appendages of of the hindwings of the female specimen, fi male simple, clearly bisegmented (coxopod- is present (for earlier authors this was a di- ite and harpago). agnostic character of Agraylea), while it is absent in the opposite hindwing ("character Wormaldia praecursor, new species ofAllotrichia"). It is difficult to discern affin- Figures 9-13, 17 ities based on male genitalia, whereas the TYPE: Male holotype in a piece of yellow structure of the female last abdominal seg- amber from Sayreville, Middlesex Co., New ments is almost certainly like present-day sg. Jersey, coll. Aug.-Nov. 1993, Case, Leggett, Allotrichia. The existing paleontological ev- and Borodin; AMNH NJ-87A. 1 995 BOTOSANEANU: TRICHOPTERA FROM TURONIAN AMBER 5

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Figs. 9-13. Wormaldia praecursor, n. sp., male holotype. 9. Maxillary palpus. 10. Distal parts of left forewing. 11-12. Genitalia, lateral, under very slightly different angles. 13. Inferior appendages in ventral view.

DESCRIPTON: Forewing length 4 mm. Head median cell extended more distal than dis- very opaque (ocelli, for instance, cannot be coidal cell, but bases of the two cells at the seen), but palpi distinct. Maxillary palpi with same level; fi_5 present (fi, f2, and f4 sessile; first two segments very short, segments 3 and f3 relatively slightly petiolate). No relevant 4 much longer and of about the same length, details visible in hind wing. Appendage on segment 5 extremely long (longer than all oth- sternite VII cannot be seen. Fine details of ers together). Spurs 2, 4, 4. Venation of left genitalia not observable, but general struc- forewing not entirely distinct, but what can ture easily observed in lateral view: superior be seen (fig. 10) is extremely clear, leaving no appendages very slender, slightly sinuous, doubt about this species being a typical phil- pointed, setose; segment X stout, rather high, opotamoid; discoidal and median cells closed, without a distinct point; phallus can be seen; 6 AMERICAN MUSEUM NOVITATES NO. 3140

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14

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Figs. 14-17. Photomicrographs of New Jersey amber Trichoptera. 14. Entire piece of amber with male and female of Agraylea (Nanoagraylea, n. sg.) cretaria, n. sp. 15. Habitus of male allotype. 16. Habitus of female holotype. 17. Wormaldia praecursor, n. sp. inferior appendages reaching beyond all other Lower Cretaceous of Mongolia); Dajella Su- parts ofthe genitalia, parallel-sided, apex ob- katsheva, 1990 (genus extinct, one species tuse (like a pointed arch at certain angles); a from Mesozoic deposits of the Tchita region biarticulation of the inferior appendages is of southern Siberia); Arkharia Sukatsheva, merely surmised, especially because of a su- 1982 (genus extinct, one species from the Up- perimposed leg, but I have no doubts about per Cretaceous-Maastrichtian/Danian ofthe its existence (2nd joint short). Amur region, Far Eastern Russia); two pos- ETYMoLoGy: Praecursor (L.) = forerunner. sible representatives ofthe family, not named, DIscussIoN: At present there are 21 species are known from the Upper Cretaceous (Tu- described, or at least mentioned, from fossil ronian) ofKazakhstan and respectively from material, belonging to this family. It should the Upper Cretaceous amber (Turonian- be mentioned that there are obscure or con- Coniacian?) ofTaimyr Peninsula; Dolophilus troversial generic and even familial attribu- McLachlan, 1868 (presently considered a tions of some of these species. Fossil philo- synonym of the surviving genus Wormaldia potamids were described in the following McLachlan, 1865, but for one of the fossil genera, here more or less chronologically ar- species the genus Ulmerodina Ross, 1956, was ranged: Prophilopotamus Sukatsheva, 1973 created; five species, all extinct, were de- (genus extinct, one species from the scribed in Dolophilus, one from the Upper of Kirgisia); Baga Sukatsheva, 1992 (genus Cretaceous amber of Tennessee, four from extinct, two species from the Middle Baltic Oligocene amber; and two extinct spe- and, respectively, from the Upper Jurassic- cies described in Wormaldia, from Saxonian 1995 BOTOSANEANU: TRICHOPTERA FROM TURONIAN AMBER 7

Miocene amber); Philopotamus Stephens, differs from all five by the proportions ofthe 1829 (genus surviving, one species from Bal- articles of the maxillary palpi, the venation tic Oligocene amber); Electracanthinus Ul- of the forewings, and genitalic characters. It mer, 1912 (genus extinct, one species from is quite possible that "Dolophilus (?) prae- Baltic Oligocene amber); Chimarra Stephens, missus" Cockerell, 1916, from Upper Cre- 1829 (genus surviving, five species from Do- taceous amber from Tennessee also belongs minican amber, lower Miocene/upper Oli- to Wormaldia. Dolophilus praemissus was, gocene). I could not find the description of up to now, the only trichopteran described Archiphilopotamus Sukatsheva, mentioned from North American amber. It was de- in the bibliography from the Upper Jurassic scribed from only a forewing (Cockerell, of Siberia, Krasnoiarsk region. 1916). Wormaldia praecursor, n. sp., is the oldest known species definitely belonging to the ge- Additional Note nus Wormaldia. In this multiform genus with tens of recent species, there are also three In the amber piece containing the speci- species from Oligocene Baltic amber: W. ae- mens ofAgraylea, there is also a pair ofwings qualis (Hagen, 1856), W. media (Ulmer, of a very small ; they are, unfortu- 1912), and W. congener (Ulmer, 1912). In nately, almost entirely superimposed, and it Ulmer's monograph (1912) these species were is impossible to make out the venation in a put under Dolophilus, a synonym of Wor- reliable manner. The very small size, and the maldia. Two other species of the genus were fact that the vein SC in the hindwing seems described (Mey, 1986) from Saxonian amber to be blackened, point to Hydroptilidae; the (Miocene, ca. 22 Ma): W. angularia Mey, hindwing is, nevertheless, too broad and its 1986 and W. contiguaMey, 1986 -both con- tip too blunt, somehow reminiscent of the sidered to be in the same species group as the hindwing of Lype (Psychomyiidae). Baltic amber species. The new species clearly

REFERENCES Botosaneanu, L. New Jersey. Am. Mus. Novitates 2949: 1981. On a false and a genuine Caddis-Fly from 27 pp. Burmese amber (Insecta: Trichoptera, Marshall, J. E. Homoptera). Bull. Zool. Mus. Univ. 1979. A review of the genera of the Hydrop- Amsterdam 8(10): 73-78. tilidae (Trichoptera). Bull. Br. Mus. Nat. 1992. Trichoptera of the Levant-Imagines. Hist. (Entomol.) 39(3): 135-239. Fauna Palaestina, Insecta VI. Jerusa- Mey, W. lem: The Israel Acad. of Sciences and 1986. Die K6cherfliegen des Sachsischen Humanities, 291 pp. Bernsteins (II). Dtsch. Entomol. Z.N.F. Cockerell, T. D. A. 33(3-5): 241-248. 1916. Some American fossil . Proc. U.S. Ulmer, G. Nat. Mus. 51(2146): 89-106. 1912. Die Trichopteren des Baltischen Bern- Grimaldi, D. A., C. W. Beck, and J. J. Boon steins. Schr. Phys.-okon. Ges. Kon- 1989. Occurrence, chemical characteristics and ingsb., Beitr. Naturkd. Preussens 10: 1- paleontology of the fossil resins from 380. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum ofNatural 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]

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